Donavl N A. LECKENBY DIANE M. CONCANNON

Home , Owyhee River, Pedicularis attollens, Polyctenium

GENERAL TECHNICAL REPORT PNW-120 1981 , EDITOR’S

PLANT COMMUNITIES AND THEIR IMPORTANCE TO WILDLIFE

This file was created by scanning the printed publication. Mis-scans identified by the software J. EDWARD DEALY have been corrected; however, some errors may remain. DONAVl N A. LECKENBY a DIANE M. CONCANNON

PACIFIC NORTHWEST FOREST AND RANGE EXPERIMENT STATION FOREST SERVICE U. S. DEPARTMENT OF AGRICULTURE \ ABSTRACT Plant communities in the Great Basin of southeastern Oregon are described, and a field key is provided. The value of a plant community’s vertical and horizontal structure and the seasonal availability of its forage are examined in relation to wildlife habitat in managed rangelands. Further, the importance of individual and combined plant communities to wildlife in managed rangelands is discussed, and management alternatives are presented. KEYWORDS: Communities (plant), range management, wildlife habitat.

THE AUTHORS J. EDWARD DEALY is Principal Research Plant Ecologist, USDA Forest Serv- ice, Pacific Northwest Forest and Range Experiment Station, La Grande, Oregon. DONAVIN A. LECKENBY is Research Wild- life Biologist, Oregon Department of Fish and Wildlife, La Grande, Oregon. At the time this research was conducted, DIANE M. CONCANNON was Range Conservationist, U.S. Department of the Interior, Bureau of Land Management, Vale, Oregon. This publication is part of the series pendent treatment of a specific subject, when Wildlife Habitats in Managed Rangelands - combined in sequence, the individual parts will The Great Basin of Southeastern Oregon. The be as chapters in a book. purpose of the series is to provide a range Individual parts will be printed as they manager with the necessary information on become available. In this way the information wildlife and its relationship to habitat condi- ’ will be more quickly available to potential tions in managed rangelands in order that the users. This means, however, that the sequence manager may make fully informed decisions. of printing will not be in the same order as the The information in this series is specific to final organization of the separates into a com- the Great Basin of southeastern Oregon and is prehensive whole. generally applicable to the shrub-steppe areas A list of the publications in the series, their of the Western United States. The principles current availability, and their final organiza- and processes described, however, are gener- tion is shown on the inside back cover of this ally applicable to all managed rangelands. The publica tion. purpose of the series is to provide specific information for a particular area but in doing so Wildlife Habitats in Managed Rangelands to develop a process for considering the welfare - The Great Basin of Southeastern Oregon of wildlife when range management decisions is a cooperative effort of the USDA Forest are made. Service, Pacific Northwest Forest and Range Experiment Station, and United States The series is composed of 14 separate Department of the Interior, Bureau of Land publications designed to form a comprehensive Management. whole. Although each part will be an inde- Introduction The terms “stand” and “cornmunity” are used as suggested by international agreements Our purpose is to describe the major plant on terminology (Kuchler 1964, Mueller- communities and examine the importance of Dombois and Ellenberg 1974). The specific their structure and species composition to unit of vegetation observed in the field is called wildlife. Plant communities here are directly a vegetation stand or stand (coenosis). The related to the wildlife covered .in the other generalized, abstract unit of vegetation chapters and can be used in developing analogous to an “average” of a group of management plans based on characteristics similar stands is a plant community or vegeta- and potential of the plant community. We did tion type. A plant community with a “definite not cover some northern Great Basin flora, uniform habitat, and physiognomy” is rangeland communities because they are rare, called an association (Mueller-Dombois and small in total area, or information is not Ellenberg 1974, p. 174). available. Plant communities presented here are com- Great Basin rangelands in southeastern pared with Kkhler’s (1964) vegetation types Oregon support a wide variety of plant com- in table 1. Kuchler has one major vegetation munities, dominated by grasses, shrubs, or type for each listing, which represents the trees ranging from conifers to deciduous and major physiognomy (general aspect and visual evergreen hardwoods. Predominant are big character) of a region. Kuchler omits other sagebrush communities. Tree-domina ted and similar vegetation types less extensive in true grassland communities constitute the occurrence and less noticeable or refers to them least common types. True grasslands occur as as inclusions because his treatment of vegeta- meadows, relict stands of valley bottom bunch- tion throughout the conterminous United grass, and subalpine bunchgrass types. States is so extensive. Table 1 shows.,-single Tree-dominated communities occur primarily vegetation types of Kuchler’s related to a at elevations above the sagebrush steppe, ex- number of vegetation types used in this cept willow and cottonwood communities of publication. The best example is Kuchler’s low-elevation riparian zones. Quaking aspen is (1964) “sagebrush steppe”-the big sagebrushbearded bluebunch wheatgrass vegetation primarily restricted to mountain riparian type. We recognize 11 vegetation types zones, such as streams, seeps, springs, ponds, dominated by sagebrush within Kuchler’s or lakes (fig. 1). With the exception of relict (1964) “sagebrush steppe.” The Wyoming big stands of ponderosa pine (Packard 1972) or white fir (Hansen 1956), curlleaf mountain- sagebrush/bunchgrass vegetation type mahogany requires the most moisture of the described here is probably the only one relating dry-land tree types in the high desert moun- directly to it since it is considered the most tains. Western juniper grows just below common sagebrush of the high desert in curlleaf mountainmahogany and mixes with it Oregon (Winward 1980). in transition zones. These tree-dominated com- Subtle variations in an environment give munities are adjacent to and above the shrub rise to different subordinate plant species in zone. communities. These “characteristic” and “dif- Big sagebrush, including several sub- ferentiating” species (Mueller-Dombois and species, is dominant among the shrub com- Ellenberg 1974) are useful indicators of site munities. Other significant tall shrub potential. For example, the mountain big communities include black greasewood, squaw sagebrush/bunchgrass community encom- apple, Bolander silver sagebrush, and moun- passes stands containing either Thurber tain silver sagebrush. Short shrub communi- needlegrass, western needlegrass, or needleandthread. These grasses are generally subor- ties include shadscale saltbush, and stiff, low, dinate to bearded bluebunch wheatgrass and early low, black, and cleftleaf sagebrushes. reflect an increasing sandiness of the soil pro- ‘Common and scientific names and plant symbols are from file. Recognition of these three variations Garrison et al. (1976)and are listed in the appendix.

1 Figure 1. -Elevation and site relationships among dominant plant species in the Great Basin of southeastern Oregon. Plant symbols are from Garrison et al. (1976). AGSP = Agropyron spicatum; ARAR = Artemisia arbuscula; ARCA = Artemisia cana; ARLO2 = Artemisia longiloba; ARN02 = Artemisia nova; ARRl = Artemisia rigida; ARSP = Artemisia spinescens; ARTR2 = Artemisia tripartita; ARTRT = Artemisia tridentata subsp. tridentata; ARTRV = Artemisia tridentata subsp. vaseyana; ATCO = Atriplex confertifolia; CARU = Calamagrostis rubescens; CELE = Cercocarpus ledifolius; CEVE = Ceanothus velutinus; DlSTl = Distichlis sp.; ELCl = Elymus cinereus; FElD = Festuca idahoensis; GRSP = Grayia spinosa; JUOC = Juniperus occidentalis; MURl = Muhlenbergia richardsonis; PIP0 = Pinus ponderosa; POSA3 = Poa sandbergii; POTR = Populus tremuloides; PREM = Prunus emarginata; SAVE2 = Sarcobatus vermiculatus; SlHY = Sitanion h yslrix; SYO R = Symphoricarpus oreophilus. Table l-Comparison of the plant communities with potential natural vegetation by Kuchler (1964)

Natural vegetation as described by Kuchler (1964) Plant communities in this publication 37 mountainmahogany-oak scrub Curlleaf mountainmahoganylmountain big sagebrushmunchgrass (similar position-not equal) Curlleaf mountainmahoganylmountain snowberrylgrass Curlleaf mountainmahoganylpinegrass Curlleaf mountainmahoganylIdaho fescue Curlleaf mountainmahoganybearded bluebunch wheatgrass-Idaho fescue

No provision Squaw applebNnchgrass

55 sagebrush steppe Western juniperbig sagebrushbearded bluebunch wheatgrass (withjuniper) Western juniperbig SagebrusWIdaho fescue

55 sagebrush steppe Basin big sagebrushhunchgrass Wyoming big sagebrushbunchgrass Mountain big sagebrushmunchgrass Threetip sagebrushbunchgrass Bolander silver sagebrushbunchgrass Mountain silver sagebrushbunchgrass Stiff sagebrushmunchgrass Low sagebrushhunchgrass Cleftleaf sagebrushbunchgrass Early low sagebrushbunchgrass I Black sagebrushbunchgrass

40 saltbush-greasewood Black greasewoodlgrass Shadscale saltbushbunchgrass

No provision Riparian

No provision Permanently wet meadows Seasonally wet meadows

No provision Quaking aspenlmountain big sagebrush Quaking aspenlgrass

No provision Subalpine big sagebrushbunchgrass

52 alpine meadows and barren Subalpine bunchgrass (similar-notequal)

,within the community should help managers of wildlife and livestock. The associated changes in soil texture2 and plant phenology are important to programs designed to maintain suitable habitat for animals and to increase production of forage.

%oil nomenclature and definitions are from “Soil Survey Manual” (Soil Survey Staff 1951).

3 PLANT AND COMMUNITY STRUCTURE Most animal species are more narrowly adapted to plant structure for thermal and hiding cover than they are to plant species for food. For example, mule deer and black-tailed jackrabbits use a wide variety of forage species (Hill 1956, Ingles 1967, Kufeld et al. 1973);but a specific structure of trees or shrubs (fig. 2) is required to fulfill their thermal or hiding cover requirements (Anthony 1928, Thomas et al. 1979a). The sage sparrow has seed and insect food available in many plant communities, but a specific shrub structure is required to provide hiding cover and suitable nesting sites (Gabrielson and Jewett 1940, Miller 1968). The overhanging or umbrellalike structure of bunchgrass provides both hiding and thermal cover for birds, such as the lark sparrow in the open bunchgrass communities (fig. 3).

Figure 3.-A, arrow at the base of a bunchgrass points to the nest of a lark sparrow. B, the nest is hidden from predators and is Figure 2.-Big sagebrush provides impor- protected from sun and wind in an open tant thermal cover for small mammals stand of grass by the overhanging blades (Robert R. Kindschy photograph). of grass (Chris Maser photograph).

4 - Shrub structure is variously altered from depending on type and intensity of use (Hor- browsing by different animals and at different may 1943). This species also occurs throughout intensities (fig. 4). Antelope bitterbrush, for its range in different structural forms because instance, when severely browsed produces a of- what appear to be genetically distinct crown that appears to be structurally different, populations (Alderfer 1976, Nord 1965). A

Figure 4.-A, lightly used antelope bitterbrush illustrating the open form. B, antelope bitterbrush heavily browsed by deer illustrates the dense, hedged form. C, antelope bitterbrush heavily browsed by cattle develops a low, dense, mushroom- shaped crown that hampers nesting of some bird species within the crown but provides good thermal and hiding cover for a variety of vertebrate species.

5 The structural diversity of rangeland com- stands is on a scale too small to meet the home munities relates directly to wildlife diversity: range needs of a species that requires large the greater the structural diversity, the blocks of uniform vegetation (Thomas et al. greater the wildlife diversity (fig. 5). Excep- 1978, 1979~). tions may occur where a mosaic of diverse

Figure fi.-Cliffs, talus, sagebrush, juniper, seasonally and permanently wet meadows, and riparian vegetation provide habitat for a wide variety of wildlife in the western edge of the Great Basin.

6 Plant communities contain both vertical rangelands. MacArthur and MacArthur (1961) and horizontal structural elements. At least considered vertical layers of vegetation critical three vertical vegetative layers-herbaceous, to diversity of avian species. Thomas et al. shrub, and tree (fig. 6)-are provided by (1977)examined each 5-foot layer of vegetation western juniper communities in high desert from shrubs to the tops of trees and found that

Figure 6.-This tree=shrub=grasscommunity illustrates the structural variety present and the high vertical diversity in some communities of the high desert. Plant symbols are from Garrison et al. (1976). AGSP = Agropyron spicatum; ARAB1 = Arabis sp.; ARTRV = Artemisia tridentata subsp. vaseyana; ASTRA = Astragalus sp; CH NA = Chrysothamnus nauseosus; FElD = Festuca idahoensis; JUOC = Juniperus occidentalis; SlHY = Sitanion hystrix; STIPA = Stipa sp.

7 different species of birds were significantly cor- size provides a more suitable environment than related with different vertical layers in both large uniform areas of seeded grasslands or coniferous and deciduous vegetation. Horizon- dense shrublands. Tree communities, such as tal diversity, on the other hand, deals pri- western juniper and curlleaf mountain- marily with stands of different plant com- mahogany, provide valuable thermal and munities or different stages within plant hiding cover for wild ungulates (deer, prong- communities that are close together (fig. 7). horn, bighorn sheep) and other species of For example, optimum spacing between stands wildlife, as well as important shading areas for of big sagebrush and crested wheatgrass for livestock in the summer. black-tailed jackrabbits requires that the These communities with an open tree wheatgrass openings be no larger than 600 cover, besides being good wildlife habitat, pro- meters (1,900 ft) across (Westoby and Wagner duce an ideal opportunity for livestock to 1973), because most use of wheatgrass occurs obtain much needed shade close to forage. within 300 meters (985 ft) of the type edge. McIlvain and §hoop (1971) studied the use of This illustrates how the size of two adjacent shade by cattle in the southern Great Plains communities can result in optimum habitat for and found that summer-long gains by yearling a particular animal species. Hereford steers increased 9 kilograms (19 Ib) Rangelands of the desert basins, uplands, per head when shade was available. They also and mountains are limited in their complement found that shade was nearly as effective as the and acreage of tree communities and true location of water and supplemental feed in (climax) grasslands. The abundant shrub com- distributing use by cattle. Advantages of munities are predominantly big sagebrush. In shade for livestock, then, are threefold: in- the enhancement of both livestock range and creased weight gain, increased forage supply, wildlife habitat, a mixture of communities in and more uniform range conditions. These proper juxtaposition and of optimum stand advantages also apply to ungulate wildlife.

Figure 7.-A variety of habitats produces a variety of wildlife. A mosaic of plant communities of appropriate sizes and shapes maximizes edge and provides a wide selection of forage and COWQB species.

8 AVAl LAB1LlTY 0 F FORAGE ungulates to reach green regrowth that otherwise would be unavailable to them. Also, Forage availability of high desert plant important winter foraging areas of wild communities varies annually and seasonally ungulates can be enhanced by preventing live- (fig. 8). Annual herbaceous species are stock from grazing after fall rains have available as green forage in early spring but produced herbaceous regrowth. dry rapidly and provide only dry material by late spring. Most perennial herbaceous species are generally available from midspring to mid- PLANT COMMUNITY GROUPS summer, whereas deciduous shrub and tree The array of plant communities listed in species usually provide their most nutritious the following key and later described in detail forage from midspring through late summer. can be dealt with individually in management. Evergreen and deciduous shrub and tree forage Depending on the user’s objective, however, is available throughout the year, although simplified groupings of these communities nutritional value peaks in the early growing may be made (table 2). A criterion for grouping season. Annual and perennial herbaceous might be that the combined communities have species produce nutritious regrowth after fall equivalent values for livestock or for a featured rains. Although production varies among wildlife species; or the communities in the years, regrowth is usually available through- group would react in an equivalent way to a out the winter, particularly on south-facing specific management program (Leckenby exposures. 1978a). The rationale for grouping plant communities and examples of the relationships of Controlled seasonal use is the key to co- terrestrial vertebrates to such groups will be ordinated use of the same range by livestock and wild ungulates. Controlled livestock use of presented in another chapter in this series, perennial grasses can produce a stubble height “The Relationship of Terrestrial Vertebrates of some plants short enough to allow wild to the Plant Communities.’’

I DECIDUoU5 woow I I

Figure 8.-Completely shaded blocks indicate the season of greatest productivity and nutritional value for forage plants. Partially shaded blocks with arrows indicate nutritious fall regrowth.

9 Table 2-Plant communities grouped by similarity of structure and selected structural and site characteristics1

Riparian X X X X X All Highly variable in soil depth and stoniness

Moderately deep to Quaking aspen/grass deep Moderately deep to Quaking aspentmountain big sagebrush deep Shallow to moderately Curlleaf mountainmahoganyt deep; stoniness mountain big sagebrush variable Curlleaf mountainmahoganyl mountain snowberry Moderately deep; Curlleaf mountainrnahoganyl stoniness variable Idaho fescue Curlleaf mountainmahoganyl bearded bluebunch wheatgrassl Idaho fescue Curlleaf mountainmahoganyl Moderately shallow; pinegrass stoniness highly variable

Western juniperbig sagebrushl Shallow to moderately bearded bluebunch wheatgrass deep; stoniness variable Western juniperbig sagebrush1 Idaho fescue

Basin big sagebrushlbunchgrass X X Below Moderately deep to 2 140 (7,000) deep; stoniness Mountain big sagebrushl 1 070- (3,500- variable. Silver bunchgrass 2 900 9.500) sagebrush flooded in Subalpine big sagebrush1 1 070- (3.500- spring bunchgrass 2 900 9,500) Wyoming big sagebrushl Below bunchgrass 1 980 (6.500) Threetipped sagebrushl 915- (3,000- bunchgrass 1 830 6,000) 1 680- (5.500- Bolander slver sagebrush1 2 440 8,000) bunchgrass 1 680- (5,500- Mountain silver sagebrush1 2 440 8,000) bunchgrass Table 2-Plant communities grouped by similarity of structure and selected structural and site characteristics1 (Continued)

Stiff sagebrushlbunchgrass Very shallow and mostly very stony over a layer restricting Low sagebrushlbunchgrass root growth; saturated in spring Cleftleaf sagebrushlbunchgrass

Black sagebrushlbunchgrass

Early low sagebrush1 bunchgrass

Squaw applebunchgrass Moderately deep to deep

Black greasewoodlbunchgrass Deep; flooded in spring

Shadscale saltbushbunchgrass Moderately deep; no apparent layer restricting root growth

Meadow. seasonally wet X X X All Deep; saturated to flooded in sp-~g.dry in summer and fall

Meadow, permanently wet Deep; saturated to flooded in spring; perennially wet

Subalpine bunchgrass Shallow to deep; stoniness variable

IX=present Very shallow-less than 25 centimeters (10 inches); shallow-26 to 50 centimeters (10 to 20 inches); moderately deep-51 to 125 centimeters (20-49 inches); and deep-more than 126 centimeters (49 inches). Key to Plant Communities

CONSTRUCTION OF KEY The plant communities are identified in a dichotomous key by descriptive vegetation and site characteristics easily recognized in the field. Groups of communities are separated on broad characteristics, such as trees vs. no trees, broadleaf trees vs. coniferous trees, shrubs vs. no shrubs, and so forth. Here we consider curlleaf mountainmahogany a tree. Site information, such as elevation or topography, is used as an aid in distinguishing communities or groups of communities. Sep- aration within community groups may be based on vegetative differences among dominant species, such as anatomical variations in leaves alone or in combination with differences in form or height. Some native communities are not covered. Some stands will not key to a community because of their early stage of succession. In stands where community identification is difficult, users should refer to the section, “successional vegetation. ”

KEY The general riparian zone will be identified here rather than within the structure of the key because there is insufficient information from which to develop a multiple community key section: It is a zone with communities immediately adjacent to wattr, that require large amounts of free or unbound water, that may be dominated by trees, shrubs, or grasses, and that may be multilayered - (RIPARIAN ZONE COMMUNITIES, page 17).

12 Page la Communities with tree cover (tree communities not in key; see “SUCCESSIONAL . VEGETATION” page 55) 2a Cover dominated by broadleaf trees 3a Tree cover dominated by quaking aspen 4a Shrub cover of mountain big sagebrush common to abundant (QUAKING ASPEN/MOUNTAIN BIG SAGEBRUSH) ...... 18 4b Shrub cover sparse to absent (QUAKING ASPEN/GRASS) ...... 17 3b Tree cover dominated by curlleaf mountainmahogany 5a Shrub cover common 6a Shrubs primarily mountain big sagebrush (CURLLEAF MOUNTAINMAHOGANY/MOUNTAIN BIG SAGEBRUSH/BUNCHGRASS) ...... 20 6b Shrubs primarily mountain snowberry (CURLLEAF MOUNTAINMAHOGANY/MOUNTAIN SNOWBERRY/GRASS) ...... 22 5b Shrub cover sparse to absent 7a Understory primarily bunchgrass 8a Grass layer obviously dominated by Idaho fescue (CURLLEAF MOUNTAINMAHOGANY/IDAHO FESCUE)...... 22 8b Grass layer not obviously dominated by Idaho fescue; grass a codominant mixture of bearded bluebunch wheatgrass and Idaho fescue (CURLLEAF MOUNTAINMAHOGANY/BEARDED BLUEBUNCH WHEATGRASS-IDAHO FESCUE) ... 22 7b Understory not primarily bunchgrass; grass predominantly pinegrass (CURLLEAF MOUNTAINMAHOGANY/PINEGRASS) ... 24 2b Cover dominated by coniferous trees; dominant tree western juniper with a shrub layer dominated by big sagebrush 9a Grass cover dominantly bearded bluebunch wheatgrass (WESTERN JUNIPERlBIG SAGEBRUSH/BEARDED BLUEBUNCH ” WHEATGRASS) ...... 25 9b Grass cover dominantly Idaho fescue (WESTERN JUNIPER/BIG SAGEBRUSHDDAHO FESCUE)...... 25 lb Communities without tree cover 10a Communities with shrub cover dominant; grasses and forbs mostly native (shrub communities not in key; see “SUCCESSIONAL VEGETATION” page 55) 1la, Dominant mature3 shrubs taller than 5 decimeters (20 inches) 12a Dominant shrub a sagebrush species4 13a Dominant shrub with persistent (overwintering), entire (leaf edges smooth), and lance-shaped leaves 14a Leaves of dominant shrub silver-gray, plants in internally drained basins with seasonal flooding; associated herbaceous vegetation usually sparse; soil alkaline and extremely clayey (BOLANDER SILVER SAGEBRUSHlBUNCHGRASS). ... 40 sMature sagebrush considered at least 20 years old by ‘Sagebrush and attendant site characteristics from Win- count of annual rings or xylem layers of main stem (Win- ward (1980)and Winward and Tisdale (1977). ward 1980). This should also be the guide for other shrub species.

13 Page 14b Leaves of dominant shrub green-gray, plants along margins of streams or meadows; associated herbaceous vegetation floristically rich and highly productive; variable soils (MOUNTAIN SILVER SAGEBRUSH/BUNCHGRASS).... 40 13b Dominant shrubs with persistent lobed and wedge-shaped leaves 15a Dominant shrub uneven topped, with flowering shoots (inflorescences) arising throughout the crown (bottom to top); branches appear as upright feathery sprays 16a Mature shrubs greater than 100 centimeters (40 inches) tall; leaf margins straight, forming a tapered wedge; common big sagebrush community of valley bottoms and foothills on deep, well-drained soils (BASIN BIG SAGEBRUSH/BUNCHGRASS)...... 31 16b Mature shrubs less than 100 centimeters (40 inches) tall; leaf margins flare outward, forming a bell-shaped wedge; sagebrush on dry sites commonly below 1 830 meters (6,000 ft) and on moderately deep, well-drained soils (WYOMING BIG SAGEBRUSH/BUNCHGRASS).... 37 15b Dominant shrubs even topped with flowering shoots (inflorescences) arising from the upper portion of the crown and extending above the foliage 17a Dominant shrub with leaf margins belled outward; flower stalks absent, flowers attached directly to main stem of flowering shoot (inflorescence spicate) or flower stalks present but unbranched (inflorescence racemose); widespread through the high desert steppe (LOW SAGEBRUSH/BUNCHGRASS)...... 44 17b Dominant shrub with leaf margins not belled outward, leaf margins straight; flower stalks branched (inflorescencepaniculate) 18a On dominant shrub, four to six individual (very small) flowers per head; found on deep, well-drained soils throughout the upper foothill and mountain areas where soil moisture is available most of the summer (MOUNTAIN BIG SAGEBRUSH/ BUNCHGRASS) ...... 35 18b On dominant shrub, more than six individual (very small) flowers per head; often roots from branches in contact with soil (layering);found at high elevations (most commonly above 2 000 meters or 6,500 ft) (SUBALPINE BIG SAGEBRUSH/ BUNCHGRASS) ...... 35 (Treated briefly under mountain big sagebrush/ bunchgrass community description) 12b Dominant shrub not a sagebrush species 19a Dominant shrub spiney, bark is whitish; usually found on alkaline soil where water table is shallow-flood plains, playas, and terraces (BLACK GREASEWOOD/GRASS)...... 49

14 Page 19b Dominant shrub not spiney, bark is dark; usually occurs on high moist hillsides in the northern edge of the high desert steppe; often near the ponderosa pine zone and often mixed with antelope bitterbrush (SQUAW APPLEIBUNCHGRASS)...... 48 1lb Dominant mature shrubs shorter than 5 decimeters (20 inches) 20a Dominant shrub with leaf edges smooth (leaves entire), spines present on branches; occurs primarily on alkaline soils in desert plains and foothills (SHADSCALE SALTBUSH/BUNCHGRASS) ...... 51 20b Dominant shrub with leaves divided or lobed, lacking spines 21a Leaves of dominant shrub divided into lobes; lobe length more than three times lobe width 22a Dominant shrub with flowering shoot leaves divided; flower stalks absent, flowers attached directly to main stem of flowering shoot (inflorescence spicate); all leaves winter deciduous; most commonly found in the north and west edge of the Great Basin and beyond, in Oregon, on shallow rocky soils (STIFF SAGEBRUSH/BUNCHGRASS) ...... 42 22b Dominant shrub with flowering shoot leaf edges smooth (entire) 23a Flower stalks branched (inflorescence paniculate); leaves on upper flowering shoots much longer than flower heads; found in the northern part of the Great Basin in Oregon and north; grows on moderate to deep, well-drained, loamy and sandy loam soils (THREE-TIP SAGEBRUSH/BUNCHGRASS)...... 38 23b Flowering shoots with flower stalks (pedicels) unbranched (racemose) or absent (spicate); flowering shoot leaves equal to or only slightly longer than flower head; most common to the north of the Great Basin in Oregon, occurring only at moderately high elevations of 1 500 to 2 700 meters (5,000 to 9,000 ft), and usually within the conifer zone (CLEFTLEAF SAGEBRUSH/BUNCHGRASS)...... 44 (Cleftleaf sagebrush treated briefly under low sagebrus Wbunchgrass) 21b Leaves of dominant shrub divided into lobes; lobe length less than

three times lobe width / 24a Flowering shoots (inflorescences) of dominant shrub with flower stalks (pedicels) branched (paniculate); fruits mature after August; seed stalks brownish and persistent into following year; leaves very small; stands of plants appear dark in color compared with low sagebrush; occurs on droughty, stony, and often calcareous soils (BLACK SAGEBRUSHIBUNCHGRASS) ...... 47 24b Flower shoots (inflorescences)of dominant shrub with flower stalks unbranched (racemose) or absent (spicate); seed stalks grayish and weakly persistent into following year 25a Fruits of dominant shrub mature in July or early August (EARLY LOW SAGEBRUSH/BUNCHGRASS)...... 47

15 Page 25b Fruits of dominant shrub mature from late August to October; widespread throughout the Great Basin of southeas tern Oregon (LOW SAGEBRUSH/BUNCHGRASS)...... 44 10b Communities without shrub cover, shrub cover sparse, or shrub cover sparse with introduced grass species (grass or forb communities not in key; see “SUCCES- SIONAL VEGETATION,” page 55) 26a Communities reseeded with introduced grasses and with or without introduced forbs, or communities appearing as bunchgrass types below 2 440 meters (8,000 ft) with or without a sparse shrub cover; mostly sagebrush communities with the shrub component eliminated or reduced by fire, chemicals, or mechanical means 27a Rehabilitated with introduced species; old seedings may have big sagebrush or rabbitbrush, and native grasses and forbs mixed at various densities with crested wheatgrass; seedings sometimes include introduced forbs (CRESTED WHEATGRASS SEEDINGS-not discussed in text) 27b Rehabilitated without introduced species (shrub cover removed by fire, chemical, or mechanical means); residual native species used for natural reseeding of grass stand; often dominated by bearded bluebunch wheatgrass, with a mix of other species present (SUCCESSIONAL VEGETATION)...... 55 26b Communities not reseeded 28a Sod-forming grasslands 29a Occurring in permanently wet situations along streams, ponds, or lake (PERMANENTLYWET MEADOWS) ...... 54 29b Occurring in places that are seasonally wet a significant part of the year; near streams or near other moist areas (SEASONALLY WET MEADOWS) ...... 52 28b Not sod-forming grasslands; bunchgrass communities in the subalpine zone above 2 440 meters (8,000 ft) (SUBALPINE BUNCHGRASS)...... 55

16 Riparian Zone Communities Management decisions must be made on a case by case basis because of the high varia- Riparian zones are identified by the bility between communities in the zone. Many presence of vegetation that requires free or un- critical site factors that influence productivity bound water or conditions that are more moist change from one riparian community to than normal (Thomas et al. 1979b). Riparian another . plant communities within this zone are complex and highly variable in structure, number of species, species composition, productivity, Quaking Aspen Communities and size. This variability is due, in part, to the interaction of water quality, hydraulics, Quaking aspen communities are scattered hydroperiod, topography, soil, geology, eleva- through mountainous areas of the Great Basin, tion, animal use, and alterations by people most commonly at elevations from 1 550 to (Odum 1971, 1979; Thomas et al. 1979b). They 2 150 meters (5,000 to 7,000 ft). These com- are disproportionately important among Great munities are usually found on wet sites but Basin plant communities, yet there is probably occasionally occur in combination with moun- less known about them than any other major tain big sagebrush. zone under management. Because of the complexity of the riparian 2-QUAKING ASPENlGRASS (fig. 9) zone and a lack of information on riparian Vegetation vegetation in the Great Basin of southeastern Oregon, this section is limited to a general Crown cover of quaking aspen in the quak- discussion. The riparian zone in relation to ing aspedgrass community varies from 25 to management is discussed by Thomas et al. 75 percent. Some stands have a two-tiered tree (197913). canopy-a mature and a reproduction layer. Mature stands reach a mean height of 21 1-RIPARIAN meters (70 ft) with a mean stem diameter Riparian communities vary structurally from a low grass and grasslike layer to a tree layer reaching above 25 meters (80 ft) and include many combinations of grasses, forbs, shrubs, and trees in between. Density of vegetation varies markedly. Because most woody vegetation is deciduous, however, hiding and thermal cover is effective primarily during the growing season. Concentrated and valuable forage resources characterize this community. Inten- sive grazing may alter vegetation, causing not only a loss of forage but a different community structure and reduced productivity for the site. Soil losses through erosion of streambanks and lowering of water tables through channeli- zation may permanently reduce plant productivity or change the vegetation. For example, a Figure 9.-Quaking aspenlgrass community streambed lowered by erosion can cause a along an intermittent streambed. Note the lower water table in the adjacent meadow and lack of reproduction and close cropping of grass and forbs by livestock and mule convert the meadow to a shrub community. deer.

17 (breast high) of 36 centimeters (14 inches) and a Soil maximum diameter of 56 centimeters (22 Soils are moderately deep to deep, gravelly inches). Basal area (area of tree trunks cover- silt loams and loams. Cobble and stone volume ing the ground surface) of one large mature in the soil varies, and coarse fragments on the stand was 28 square meters per hectare (120 ft2 surface are scarce. These soils are well drained, per acre).s When present, the shrub layer is runoff is medium, and erosion hazard is sparse and consists of occasional plants of moderate. mountain big sagebrush, green rabbitbrush, or mountain snowberry averaging 40 centimeters (16 inches) in height. Discussion Grass and forb species, plant composition, Most stands have been intensively grazed and plant density vary among stands, partially by livestock for nearly 100 years. Since these sites are highly productive and found near because of varying amounts of moisture and severity of livestock use. Common grasses in- water, wild and domestic ungulates concentrate on them. Reproduction of aspen on clude blue wildrye, wheatgrass, needlegrass, hairgrass, or bottlebrush squirreltaiP (see foot- streamsides is damaged by grazing, except in note 5). Total crown cover of the grass-forb areas where access by livestock has been layer may vary from less than 5 to over 30 per- restricted. cent. Height of the grass layer averages about Beavers inhabit this community where 40 centimeters (16 inches). there is sufficient running water. Damage to Stinging nettle, a forb, often dominates aspen reproduction by livestock and wild disturbed stands. Other common forbs include ungulates and to older trees by beavers has peavine, aster, dandelion, sorrel, cinquefoil, eliminated some stands, thereby eliminating and bedstraw (see footnotes 5 and 6). False beaver habitat. In one area, for example, there hellebore and thistle are occasionally found. was one active beaver pond, one being built in Dominant forb species average 1 meter (3.28 ft) an aspen stand too small to support the beaver, tall. Some stands, however, lack tall forbs, and approximately 15 abandoned ponds where such as stinging nettle, sorrel, false hellebore, aspen stands have been eliminated. It did not and thistle. Height of forbs in such stands appear that these destroyed stands would become reestablished (see footnote 5). averages 30 centimeters (12 inches). Aspen stands provide valuable thermal and Site hiding cover for many vertebrate species, and Elevation of this community ranges be- nesting sites for birds, including cavity nesters. tween l 830 and 2 300 meters (6,000 and 7,500 ft) in southeastern Oregon. It may occur near the top of mountains or on the middle, lower, 3-QUAKING ASPENlMOUNTAlN BIG toe, and bottom slopes; it often occurs around SAGEBRUSH (fig. 10) springs, seeps, lakes, ponds, or along streams Vegetation on slopes from 0 to 30 percent. Some stands are extensive, occurring adjacent to and A single species tree stand of quaking parallel with streams in unbroken bands up to aspen is typical of this community. It usually 150 meters (500 ft) wide and several hundred occurs as an open stand with less than 50- meters long. Other stands are small patches of percent crown cover. Height ranges from 6 to less than 1 hectare (2.5 acres) in size. 12 meters (20 to 40 ft), and trees on these sites often appear stunted and broad crowned with sJ.Edward Dealy, unpublished data on file at Range and heavy lateral branches (see footnote 5). Occa- Wildlife Habitat Laboratory, La Grande, Oregon. sionally, all mature trees in this type are either sPersonal communication from A. H. Winward, Oregon dying or dead. Some stands have a dead (snag) State University, Corvallis. overstory above live reproduction, but in other

18 is a consistent but sparse component; it increases in density, however, as moisture availability increases in severely grazed communities (see footnote 5). Common perennial grasses and grasslike plants in disturbed stands of this type, in descending order of amounts of crown cover, are needlegrass, bottlebrush squirreltail, wheatgrass, and sedge. Annual cheatgrass is also common. Total crown cover of these species is less than 10 percent. Mean heights of needlegrass and wheatgrass .are 50 and 60 centimeters (20 and 24 inches), respectively. Bottlebrush squirreltail and cheatgrass both average 25 centimeters (10 inches) in height, whereas sedge averages 5 centimeters (2 inches) (see footnote 5). Forbs are sparse. Thistle and aster appear as occasional plants, and common dandelion is a rare component. Their total crown cover is generally less than 1 percent (see footnote 5).

Figure 10.-Quaking aspenlmountain big Site sagebrush community illustrating the The quaking aspenlmountain big common branched tree form. Note the sagebrush community occurs most commonly hole in the trunk used by cavity nesters on what appear to be dry sites between 1 550 (Chris Maser photograph). ' and 2 150 meters (5,000and 7,000 ft). It is present both in the Owyhee Uplands and Basin stands both layers are dead. The causes of such - and Range Provinces (Franklin and Dyrnes-s mortality are unknown. It does appear, how- 1973). Macrorelief is mountainous and micro- ever, that these sites are marginal, and fluctua- relief smooth. Slope ranges from less than 5 to tion in underground moisture or winter 25 percent, and the position on a slope may be snowpack may be important factors. Also, upper, middle, or lower. Slope aspects are long-term concentrated use of these small generally northerly -from nor th-nor t hwest to areas for forage and shade by livestock and big east-northeast (see footnote 5). game may have consistently destroyed reproduction and prevented replacement of the old, Soi I degenerating stand. Soils are moderately deep, gravelly loams Crown cover of quaking aspen reproduction to sandy loams overlying basalt or rhyolite. ranges from 25 to 75 percent, and all heights They are well drained, and erosion hazard is may be represented, from a few centimeters to moderate. Cobble and stone volume in the soil mature height. It is common, however, to see ranges from 5 to 30 percent, and coarse the aspen reproduction, where present, domin- fragments cover less than 5 percent of the soil ated by a single age and height class. surface (see footnote 5). Mountain big sagebrush is the dominant shrub, with crown cover ranging from 15 to 40 Discussion percent. Average height is approximately 1 This quaking aspen community indicates a meter (3.28 ft). Gray rabbitbrush is the next high soil moisture level, possibly caused by the most abundant shrub and may have crown combination of an unusual underground source cover as high as 20 percent. Green rabbitbrush of water and an exceptionally heavy snowpack.

19 All observed sites occur near maximum eleva- grass, tree-snowberry-grass, tree-bunchgrass, tions on aspects roughly 180" from prevailing and tree-grass (Dealy 1971,1975). winter storm winds. Snowpack is deep and persistent compared with surrounding sites. Use by livestock is intense in this community because of its high production of forage and tree shade. These stands are small; relative to the overall livestock forage resource of the adjacent open range, they have little impact. They appear to be important, however, as islands that maintain habitat diversity for wildlife and, therefore, merit management. Curl leaf Mountainrnahogany Communities (figs. II and 12) Curlleaf mountainmahogany has two recognized varieties in the intermountain west. Only Cercocarpus ledifolius var. ledifolius is commonly present in the Great Basin of southeastern Oregon, occurring within four broad community groups: tree-sagebrush-bunch- Figure 12.- Dense curlleaf mountainmahogany stand upslope from a wigorous mountain big sagebrush stand. These two high value habitats commonly occur adjacent to one another and, as a result, increase diversity fo? deer (in foreground) and other wildlife. Precipitation is above 50 centimeters (20 inches).

4-CURLLEAF MOUNTAIMMAHOGANY/ MOUNTAIN BIG SAGEBRUSH/ BUNCHGRASS (fig.13) Vegetation This community is characterized by a patchy distribution of curlleaf mountainmahogany with a crown cover ranging from 35 to 60 percent and averaging 50 percent. Tree density varies between 800 and 2,000 stems per hectare (300 and 800 per acre) and averages 1,250 per hectare (500 per acre) with an occa- Figure I1. -Extensive stands of curlleaf sional interspersed western juniper. Occa- mountainmahogany blanket the tops and sionally, ponderosa pine may occur within the slopes of high desert mountains. Coma stands. munities with open stands and understories of mountain big sagebrush differ The shrubby understory usually consists of from those with dense-canopied stands a mountain big sagebrush layer of less than 5- and a grass understory. Vegetation percent crown cover. Density and vigor of changes depend on changes in aspect and sagebrush increase noticeably in the spaces elevation which produce changes in between trees. Gray rabbitbrush, green rabbit- available moisture. brush, wax currant, and Wyeth eriogonum are less common in the shrub layer.

20 species and some atypical species because of ecotonal influences within the narrow strip of the type.

Soi I The soil is well drained and varies from gravelly loam to gravelly sandy loam in the A horizon, and from gravelly clay loam to fine sandy loam in the B or AC horizon. Soil depth varies from 23 to 85 centimeters (9 to 33 inches) and averages 55 centimeters (22 inches). Soil surface cover of coarse fragments ranges from 1 to 60 percent and averages 32 percent. Cobble and stone volume in the soil ranges from 25 to 90 percent and averages 68 percent. Parent material can be either residuum or colluvium. Underlying material is Figure 13. -Curlleaf moun tainmahoganyl generally either basalt or rhyolite. The soil is mountain big sagebrushlbunchgrass com= often shallow in stringer sites adjacent to rim- munity in the Great Basin of southeastern rock, and stoniness of these soils is generally Oregon. Mountain big sagebrush grows pronounced (Dealy 1975). densely and with high vigor under open tree canopies. Discussion Dominant grasses are Idaho fescue, cut- This community has had a long history of ting wheatgrass, and big bluegrass. Other com- heavy use by livestock-sheep in the early mon grasses or grasslike species are California days and cattle more recently. This has brome, bottlebrush squirreltail, Lemmon resulted in an increase of understory species, needlegrass, and Ross sedge. such as gray or green rabbitbrush, cheatgrass, bottlebrush squirreltail, Lemmon stipa, or Between 20 and 30 forbs occur in the under- thistle. Generally, stands occur in a mosaic story. Lambstongue groundsel, arrowleaf bal- with mountain big sagebrush where relatively samroot, western hawkweed, lupine, violet, dense clumps of curlleaf mountaipmahogany and annual agoseris are most common. are interspersed with openings dominated by sagebrush and bunchgrass. Site Fire easily kills curlleaf mountain- The curlleaf mountainma hoganylmount ain big sagebrusWbunchgrass community occurs mahogany. In many instances, patterns of mostly on southerly facing slopes, ranging burning are indicated by old, large trees in rocky, fire-resistant sites adjacent to young from 0 to 40 percent at elevations of 1 600 to (60-80 year) stands in less rocky sites more 1 950 meters (5,250 to 6,400 ft). Mean elevation vulnerable to fire. The old trees appear to have is 1 750 meters (5,750 ft). Macrorelief is moun- been missed by fire and provided the seed to tainous and microrelief smooth. Since local reestablish adjacent stands. Many curlleaf topography in some areas is abruptly broken mountainmahogany stands appear to have in- by ridges and hillocks, directional aspects may creased in size since fire has been controlled change within short distances. (Dealy 1975). Such action has been an advan- Stands of this community occur on the tage to wildlife in general since these stands driest sites suited to curlleaf mountain- provide important hiding and thermal cover mahogany in the high desert mountains. When for deer and bighorn sheep, as well as nesting situated as narrow stringers on rimrock sites, sites for birds and ground-dwelling mammals. the stands characteristically contain fewer

21 5-CURLLEAF MOUNTAINMAHB Other stands are found on northerly MOUNTAIN SNOWBERRY/GRAS% aspects, varying from east-northeast to northwest at elevations between 1 800 and 1 900 Vegetation meters (5,900 and 6,232 ft). Slopes range from Stands of the curlleaf mountainmahoganyl 25 to 35 percent. Macrorelief is mountainous; mountain snowberrylgrass community are microrelief generally rough. Topography is characterized by a tree crown cover, ranging steep and abrupt in some stands and moderate from 50 to 85 percent and averaging 70 per- in others. cent. Such stands may vary in size from a few hectares to extensive areas with only small Soil interspaces that may be dominated by grasses Soils range from shallow to moderately or shrubs (often mountain big sagebrush). Tree deep and from a loam texture to gravelly loam. density varies from 900 to 3,200 stems per hec- Cobble and stone cover of the soil surface tare (365 to 1,300 per acre) and averages 1,770 ranges from 5 to 48 percent, and cobble and per hectare (720 per acre). An occasional stone volume in the soil ranges from 27 to 81 western juniper may occur and, rarely, quak- percent. Parent material is generally colluvium ing aspen on moister sites (Dealy 1975). or a mixture of colluvium and residuum The shrub understory commonly is a mix- material. Some soils overlay basalt; others ture of mountain snowberry and mountain big occur over rhyolite (Dealy 1975). sagebrush. Dominance may change from one to the other in different stands. Crown cover is Discussion generally less than 5 percent. Other shrubs Density, vigor, and size of shrubs appear to commonly found are green rabbitbrush, gray be either a function of available moisture or an rabbitbrush, chokecherry, and creeping interaction of available moisture and tree Oregon grape. Saskatoon serviceberry is pre- crown cover (reduction of light) on the drier sent on about 50 percent of the area sampled. sites. Under dense tree canopies, shrubs are Other less commonly occurring shrubs are bit- small, structurally weak, and scattered. In tercherry and desert rockspirea. more open stands, however, shrubs are larger, Dominant grasses and grasslike species are more vigorous, and more dense. Increased big bluegrass or Idaho fescue in stands lightly available moisture within sites favors an in- used by livestock. Ross sedge and Sandberg crease in grasses and forbs rather than shrubs. bluegrass are common in most stands of the Stands of the community are habitat for complex. In stands heavily used by livestock, mule deer and bighorn sheep. Livestock have there is an increase in bottlebrush squirreltail, also used these areas since the 1860’s. cheatgrass, and Sandberg bluegrass. Fire control has permitted development, Dominant forb species vary among sites in maintenance, and expansion of tree stands this community. Common species on dry sites within the curlleaf mount ainma hoganylmoun- are balsamroot, lupine, prairiesmoke avens, tain snowberrylgrass community (Dealy 1975). and varileaf phacelia; on moist sites-gray hawksbeard, heartleaf arnica, meadowrue, 6,7-CURLLEAF MOUNTAlNMAHOGANYl lambstongue groundsel, and Holboell IDAHO FESCUE (fig. 14) AND CURLLEAF r ockcres s. MOUNTAlNMAHOGANYlBEARDED BLUE- BUNCH W H EATG RASS-IDAH0 FESCUE Site (fig. 15) The driest stands of this community Vegetation occupy south to southwest slopes at elevations averaging 1 770 meters (5,800 ft). Slopes Tree cover varies from an average 66 per- average 30 percent. cent in stands on the moister sites to 36

22 Figure 14.-Curlleaf mountainmahoganyl Figure 15.-Curlleaf mountainmahoganyl Idaho fescue community commonly bearded bluebunch wheatgrass-Idaho occurs adjacent to the ponderosa pine fescue community commonly occurs zone on moist northerly sites. below and adjacent to the lower edge of the ponderosa pine zone. percent on drier sites. Curlleaf mountain- Forbs are scarce in the understory. mahogany is the dominant species, but Western hawkweed and phlox are most com- ponderosa pine occurs as scattered trees where mon on moist sites, and arrowleaf balsamroot the community is adjacent to the pine zone. is most common on drier sites. Occasional Western juniper occurs as a subordinate forbs are yarrow and annual agoseris. species in some stands. Density of curlleaf mountainmahogany ranges from 500 to 2,000 Site stems per hectare (200 to 800 per acre). Stands are either patchy in distribution or occur as The curlleaf mountainmahoganylIdaho narrow bands perpendicular to the slope below fescue community is found on most aspects and adjacent to the ponderosa pine zone. They from 1 600- to 1 700-meter elevation (5,250 to occasionally encircle scab flats as bands within 5,575 ft). Northerly slopes ranging from 5 to 15 a ponderosa pine forest. percent contain a dominant understory of Idaho fescue, whereas on other aspects with The shrub layer consists of occasional big slopes ranging from 25 to 60 percent, bearded sagebrush plants and rare plants of wax cur- bluebunch wheatgrass and Idaho fescue are rant, gray rabbitbrush, green rabbitbrush, or codominant. Macrorelief is mountainous, and antelope bitterbrush. microrelief varies from smooth to rough.

Grasses and grasslike species on sites indi- Soil cating highest soil moisture are Idaho fescue (a strong dominant), bearded bluebunch wheat- Soils are generally moderately deep, well- grass, bottlebrush squirreltail, prairie j une- drained loams. Cobble and stone volume in the grass, Sandberg bluegrass, California brome, A horizon ranges from 5 percent where Idaho Ross sedge, and cheatgrass. On drier sites, fescue is dominant to 30 percent where bearded understories are bearded bluebunch wheat- bluebunch wheatgrass is codominant with grass and Idaho fescue in a codominant rela- Idaho fescue. Cobble and stone volume in B tionship. Both Sandberg bluegrass and bottle- horizons is about 65 percent. Soils are brush squirreltail are more abundant, and developed from colluvium or weathered basalt. prairie junegrass and cheatgrass remain about Permeability is moderate,. runoff is estimated the same as on moist sites. as medium, and erosion hazard is moderate (Dealy 1975).

23 Discussion Use by cattle is minor in these stands because dense (70-percent crown cover), short stands of curlleaf mountainmahogany form a natural physical barrier. The trees commonly retain dead lateral branches below the live canopy. In a dense stand, these brittle but tough branches make penetration by cattle difficult. Domestic sheep travel through these thickets, however, as evidenced by wool hang- ing from low branches (Dealy 1971). Tree crown cover averages 35 percent on sites having a high percentage of bearded bluebunch wheatgrass. Cattle move readily Figure 16.-The curlleaf rnountain- within these stands to graze the understory. ma hoganylpineg rass corn m um ity has the highest potential for production sff dry land Mule deer use all sites for cover and forage; species in the Great Basin of southeastern these sites, however, are more important to Oregon. deer for cover than for forage (Dealy 1971).

8-CURLLEAF MOUNTAlNMAHOGANYl hawkweed is found consistently in low abun- PINEGRASS (fig. 16) dance, and thistle and common dandelion increase with disturbance. Common forbs on all Vegetation sites are prairiesmoke avens, annual agoseris, Cur llea f mount ainmahoga n y lpinegra s s and yarrow. stands contain only one other tree species-an The Alaska habenaria (commonly called occasional bittercherry. Crown cover ranges Alaska rein-orchid) grows in low abundance in from 62 to 96 percent and averages 80 percent. stands showing no evidence of litter dis- Density ranges from 1,400 to 4,700 stems per turbance. This plant develops a fleshy tuber hectare (570 to 1,900 per acre) and averages delicately attached just below the interface of 2,000stems per hectare (800 per acre). litter and mineral soil. It appears in stands protected from livestock and wild ungulate tramp- Shrubs are sparse; Saskatoon serviceberry ling by a dense tree thicket but disappears and creeping Oregon grape are almost always from sites where the litter-soil interface is even present. Occasional shrubs are green rabbit- slightly disturbed (Dealy 1975). brush, gray rabbitbrush, and Wyeth eriogonum. Site Pinegrass dominates the grass layer in The curlleaf mountainmahoganylpinegrass light to moderately grazed stands. Big blue- community has been identified in the Owyhee grass is abundant, and Ross sedge, bottlebrush Uplands (Dealy 1975) at elevations ranging squirreltail, and cutting wheatgrass are com- from 1 750 to 1 920 meters (5,750 to 6,300 ft). mon. Either Idaho fescue or Lemmon needle- It consistently occupies slopes facing east- grass is codominant with pinegrass in stands northeast to north. Slopes range from 10 to 40 that have been heavily grazed by livestock. percent and average 24 percent. Macrorelief is Other common grass and grasslike species in mountainous, microrelief smooth. These sites these heavily used stands are bottlebrush experience heavy snowpacks during most squirreltail, big bluegrass, and Ross sedge. winters. Measurements over a period of years Western hawkweed is the dominant forb on have established annual precipitation at 56 relatively undisturbed sites. On disturbed centimeters (22 inches) (Dealy 1975). sites, Nuttall violet dominates, western

24 Soil Soils are moderately deep (48 centimeters or 19 inches), varying from gravelly loams to gravelly sandy loams, overlying fractured rhyolitic bedrock. Parent material is residual. Cobble and stone cover of the soil surface ranges from 0 to 10 percent and averages 3 percent; cobble and stone volume in the soil profile ranges from 15 to 80 percent and averages 44 percent. Soils under this community are well drained and moderately permeable. Runoff is medium and erosion hazard moderate to high (Dealy 1975).

Discussion Domestic sheep and cattle grazing in the Owyhee Uplands, where this community occurs, was moderate to heavy during the late 1800's and early 1900's (Griffiths 1902). Since 1934, cattle have been the primary domestic animals grazing in the area. Historically, bighorn sheep and mule deer inhabited the area. Bighorn sheep were eliminated, however, Figure 17.-Generalized distribution of near the turn of the century and did not return western juniper (shaded portion) which occurs only in the Northwest (from Dealy until 1965, when the Oregon Department of et al. 1978a). Tree densities vary within and Fish and Wildlife reestablished them in some among the different localities. mountainous areas. Mule deer have been present throughout recorded history. 9,lO-WESTERN JUNlPERlBlG The curlleaf mountainmahoganylpinegrass SAGEBRUSHlBEARDED BLUEBUNCH community exhibits what may be the highest WHEATGRASS (fig. 18) AND WESTERN potential for production of dry land species in JUNlPERlBlG SAGEBRUSHllDAHO FESCUE the high desert mountains of the northern Vegetation Great Basin. Stands of this community where the Alaska habenaria occurs appear to be Western juniper in stands of these com- closer to pristine conditions than any others. munities occurs primarily as a single overstory This community is particularly valuable to species with wide spacing. Some stands occur birds and small ground-dwelling vertebrates. with a mature overstory, whereas others have recently developed with a young overstory on sites previously occupied by big sagebrush Western J uni per Com mu nit ies communities (fig. 19). Crown cover of western juniper is less than 35 percent in most stands.' Western juniper is unique to the inter- Big sagebrush is the dominant shrub; gray rab- mountain west. Its center of development is bitbrush and green rabbitbrush are present in the large woodland of central Oregon. It also varying amounts. On moist sites, the shrub occurs throughout southeastern Oregon, layer becomes less dominant and both western southwestern Idaho, northwestern Nevada, juniper and grass increase in importance. Occa- and northeastern California in scattered, open sionally, antelope bitterbrush and broom stands, as single trees, and rarely in more snakeweed are present (Driscoll 1964, Eckert dense, extensive stands (Dealy et al. 1978a, 1957). ~~~~~~ ~~ 197813) (fig. 17). 'Donavin A. .Leckenby, Oregon Department of Fish and Wildlife, La Grande, Oregon, unpublished data.

25 prairie j unegrass and needleandthread occur in Idaho and Oregon stands (Burkhardt and Tis- dale 1969, Roberts 1975; also see footnote 7). Forbs in the juniper community include the following: locoweed, phlox, biscuitroot, fleabane, buckwheat, pussytoes, hawksbeard, annual agoseris, rockcress, arrowleaf balsamroot, and lambstongue groundsel. Driscoll (1964) made an intensive study of western juniper communities, describing vegetation, site, and soils of nine associations and two variants in central Oregon. Three of these associations-western juniperlIdaho fescue, western juniperlbearded bluebunch wheat- Figure 18.-Western juniperlbig sagebrushlbearded bluebunch wheatgrass grass, and western juniperlbearded bluebunch community illustrating a mature stand of wheatgrass-Idaho fescue-have understories open juniper. Value as protective cover is dominated by grass. The other six have under- high for ungulates, small mammals, birds, stories with prominent big sagebrush layers, and other wildlife in the juniper and the two variants have prominent antelope communities. bitterbrush layers. Tables 3 and 4 list Driscoll’s (1964) associations and present selected information on vegetation, site, and soil. Similar communities have been studied in south-central Oregon (Adams 1975, Roberts 1975). Intensive studies on western juniper communities are lacking for southeastern Oregon. Vegetation-site relationships, including plant and community structure, can be applied from Driscoll’s (1964) work where similar conditions are found.

Site The western j uniperlbig sagebrus hlbearded bluebunch wheatgrass and western juniperlbig sagebrushlIdaho fescue communities occupy Figure 19.-Before fire control and livestock level to hilly sites and ridges and northerly grazing, this area supported a mountain slopes in mountainous areas where moisture big sage brushlbunc hgrass com m un ity. AI I levels are higher than in the sagebrush steppe. western juniper trees here have become Western juniper, as a type, occurs just below established since fire control and grazing and abuts the curlleaf mountainmahogany (reduces grass fuel) reduced the frequency of fires. zone. Precipitation ranges from approximately 25 to 38 centimeters (10 to 15 inches) per year. Dominant grasses are bearded bluebunch Macrorelief is level to mountainous, and micro- wheatgrass or Idaho fescue. On some sites relief can be rough in rimrock situations or these grasses appear as codominants. Common smooth in deeper soils. Idaho fescue dominates grasses are Thurber needlegrass, bottlebrush the understory on sites with highest moisture, squirreltail, Sandberg bluegrass, and cheat- and bearded bluebunch wheatgrass is domi- grass. Burkhardt and Tisdale (1969)also found nant on the driest sites. A more varied mixture these grasses in the western juniper type in of these grasses is found on sites of inter- southwest Idaho. Occasionally plants of mediate moisture.

26 Table 3-Number of macroplots and generalized characteristics of vegetation-soil units in the central Oregon juniper zone; reproduced from Driscoll (table 2,1964)

Associated great soil Elevational Unit S;;p,’e Aspect group range

Number -Feet lunipcrudArtfmisia/FrJtiica 5 NW. to NE. Chestnut 4,250-4,500 lunipcrus/Artrmisia/Frst uca-Lupinus 4 N. to NE. Regosol in Brown zone 4,400-4,550 luniprrus/FrJtuca 5 NW. Brown 4,100-4,300 (Purrhia variant) (2) (SE. to E.) lunipcrus/Artrmiria/~~rop~ron-Chnrnact~J 5 NW. to NE. Regosol in Brown zone 3,900-4,400 Juniprrur/Artrmiria/Agropyron 5 Level Brown 2,550-2,650 Juniprrus/Agropyron 6 E. to NE. Brown 4,1504,450 (Purrhia variant) (2) (SE.) /uniprrus/Art f miria-rurrhia 5 N. to NE. Regosol in Brown zone 4.100-4,400 /uniprrus/Ab.ropyron-FeJt tica 5 E. Regosol in Brown zone 4,2504,750 s. to sw. Brown 4,000-4,400 Juniprt ur/ArlemiJia/Asropyron-Aslraga~fiJ 6 -

Table 4-Average values of selected surface and subsoil characteristics attendant to nine associations. in the central Oregon juniper zone; reproduced from Driscoll (table 3,1964)

Horizon Basal area Bare soil Available SMS1 Organic Total Associat ion perennial surface capacity, 2- to matter, nitrogen, herbs 14-inch zone A horizon A horizon A B or AC

7.4 22.1 1.41 4.78 0.2 1 IC 6.5 30.9 1.98 1.74 .10 SI SI 4.4 33.3 1.81 3.91 .I7 1 sil, c (46.8) (2.14) (2.33) (.IO) (1) (c) 5.2 52.3 .87 1.53 .08 SI SI 6.6 41.3 2.31 1 SO .08 1 c, El 3.2 51.1 1.34 2.12 .IO scl c, cl (1) 3.1 55.0 1.54 1.05 .06 SI SI 4.1 54.7 .97 1.32 .07 SI SI 3.5 45.7 1.21 1.63 .06 IC

1 SMS, soil moisture storage. 2 The AC horizon is immediately below the A horizon in Regosols. B horizon textures were taken for the finest part of that horizon. Textural classes: cl, clay loam; I, loam; scl, sandy clay loam; sit, silt loam; c, clay; and SI, sandy loam.

SoiI respects to those described by Dealy et al. (1978a, 197813) and Driscoll(l964). Soils are highly variable in western juniper communities. Relatively young stands (100 Discussion years old or younger) have been reported on deep sandy loam soils in areas supporting the Some authors have described western juni- big sagebrush type before advent of fire con- per as an “invader” in certain big sagebrush trol (Burkhardt and Tisdale 1969, 1976). Old- communities (Anderson 1956; Burkhardt and growth stands (more than 100 years) have been Tisdale 1969, 1976). Western juniper appears found on rocky rims having shallow soils. to be a strongly dominant native species but Eckert (1957), working in southeastern historically has been kept in a subordinate role Oregon, found soils supporting western juni- on some sites because of natural fires (Dealy et per to be in the Brown and Chestnut great soil al. 1978a, 197813). Since fire control has been in groups that were derived primarily from effect and since severe livestock grazing began residuum or colluvium of basalt and rhyolite in the late 1800’s, resulting in reduced grass- origin. Some, however, developed on alluvial forb fuel for carrying fires, western juniper has fans. Soil profiles were similar in many increased its range and density in some areas

27 (fig. 19). Whether the increase of western juniper communities is an “invasion” or an “expansion” is important semantically be- Woody sagebrushes dominate the aspect cause “invasion” has a negative connotation. (physiognomy) of the high desert shrub-steppe It is more important that managers objec- plant communities. These shrubs are distrib- tively recognize values of western juniper com- uted from Canada to Mexico and from the munities and retain them, if desired; this is Pacific Northwest States to the Great Plains consistent with the strategy of preserved (Beetle 1960). diversity (Bella and Overton 1972). Aromatic gray sagebrushes appear as at The western juniper community is impor- least 13 distinctive taxa in Oregon. Among tant for livestock production because of other characteristics, habitat, height, branch- available shade, wildlife habitat (Leckenby ing form, leaf shape, chromosome number, 1977, 197813; Maser and Gashwiler 1978), chromatographic response, and fluorescence in recreational activities, and erosion protection, ultraviolet light help to identify the various im- to name a few values. A provisional list of 83 portant taxa (Beetle 1960; Hanks et al. 1971, species of birds and 23 species of mammals 1973; Stevens and McArthur 1974; Winward that use western juniper communities was 1980; Winward and Tisdale 1977). The above developed by Maser and Gashwiler (1978).The characteristics except chromosome number importance of thermal qualities of western and chromatographic response, can be used in juniper communities for mule deer has been the field for identification of taxon. Heights of documented by Leckenby (1977). Old-growth sagebrushes vary between extremes of the tall stands of western juniper provide a special basin big sagebrush and the small bud sage- habitat for cavity dwellers, such as the bushy- brush. Height of other species and subspecies tailed woodrat (fig. 20). declines between those extremes in about the following order: (1)subalpine big sagebrush, (2)mountain big sagebrush, (3)Wyoming big sagebrush, (4) mountain silver sagebrush, (5)Bolander silver sagebrush, (6)threetip sagebrush, (7) stiff sagebrush, (8)early low sagebrush, (9)low sagebrush, and (10)black sagebrush (see footnote 6, p. 18). Branching appears feathery and upright in basin big sagebrush and silver sagebrushes, tabular and compact in mountain big sagebrush, and globelike or flattened in black sagebrush (fig. 21). Leaves are predominantly wedge shaped, with three lobes (occasionally five or more) at the wide tip; silver sagebrush leaves are tapered to one tip, like lance points. Alcohol or water extracts of the leaves and other parts fluoresce characteristic shades of creamish-blue or brownish-red (table 5) in long-wave ultraviolet light (Stevens and McArthur 1974, Winward 1980, Winward and Tisdale 1977). The various sagebrushes readily cross (genetic plasticity), as evidenced by similari- ties among their morphological characteristics Figure 20.-Nests of bushy-tailed wosdrats (Beetle 1960). in a hollow western juniper stem illustrate one of the many values of old growth for At least four, and probably more, sage- wildlife habitat (Chris Maser photograph). brush taxa reach the same growth stages

28 a 6' c Figure 21 .-Three major growth forms of sagebrushes: A, feathery and upright; B, tabular and compact; and C, globular or flattened.

Table 5-Characteristic color shades of alcohol extracts of persistent leaves of 12 sagebrushes during exposure to long-wave ultraviolet light (after Winward 1980)

Sagebrush Shade of fluorescence taxa Credsh-blue Brownish-red Basin big sagebrush X Wyoming big sagebrush X Stiff sagebrush X Black sagebrush X Mountain big sagebrush X Subalpine big sagebrush X Threetip sagebrush X B olander silver sagebrush X Mountain silver sagebrush X Low sagebrush X Clef tlea f sagebrush X Early low sagebrush X

(phenology) at different times. For example, Winward and Tisdale (1977) discussed mountain big sagebrush initiates growth 2 several management reasons for identifying weeks later and ripens seed 2 weeks earlier sagebrushes to subspecies. Briefly, the various than basin or Wyoming big sagebrush; sub- taxa of sagebrush provide indicators for alpine big sagebrush initiates growth later and (1)recognition of the plant environment or site seed development earlier than any other taxon; (clim,ate,soil, and other factors); (2) estimation differences between basin and Wyoming big of productive potential; (3)prediction of reac- sagebrushes are not consistent (Winward and tion to disturbance (changes in plant composi- Tisdale 1977). tion or stand structure); (4) determination of

29 range condition; (5)evaluation of forage prefer- 363 mm in the Artemisia zone and never less ences among stands; (6)applications of grazing than 408 mm in the Pinus forest” (Daubenmire systems; and (7)selection of treatment 1956, p. 142);these moisture levels defined the methods and schedules appropriate to accom- sagebrush-forest ecotone in portions of Wash- plishment of management goals. ington and Idaho. In another example, the threetip sagebrush-fescue zone is separated Sagebrushes can be separated conveniently from a big sagebrush-wheatgrass zone and is into tall and short groups based on species, associated with cooler temperatures from subspecies, and form. Each group requires dif- October through April and with higher ferent manipulations to meet management precipitation and water balances in September, objectives for forage or cover. The tall November, December, January, and March sagebrush group is comprised of basin big (Daubenmire 1970). sagebrush, mountain big sagebrush, Wyoming big sagebrush, subalpine big sagebrush, Microclimate is closely reflected by many threetip sagebrush, mountain silver sage- plant species, and most of these are probably brush, Bolander silver sagebrush, and other as not dominants in the stand. yet undefined taxa. The short sagebrush group Grazing management has a marked influ- is comprised of low sagebrush, cleftleaf sage- ence on densities and crown cover of sagebrush, early low sagebrush, black sagebrush, brushes and cheatgrass. stiff sagebrush, bud sagebrush, and other as yet undefined or unidentified taxa. Unusual and prolonged increases in soil moisture (i.e., water spreading as a manage- The recognition of species, subspecies, and ment tool) drastically alter the relative forms of sagebrush supplies much information dominance of western juniper, sagebrushes, about a stand or a community-much that and grasses. If flooding is of sufficient dura- would take years to measure by other means; tion, pygmy woodlands and shrub steppes are for example, weather stations. Variation in converted to meadows or marshlike plant com- composition of plants among stands may iden- munities. Again, the characteristic species tify differences in environments (Fosberg and within the stand reflect the management. Not Hironaka 1964); for example, stands of silver all interactions are obvious. There is still much sagebrush, low sagebrush, and big sagebrush to learn about a site from composition of plant are usually separated by sharp ecotones species and how best to manage the site for because these species greatly differ in specific goals (Daubenmire 1968, Pechanec et tolerance to soil moisture saturation. Silver al. 1965). sagebrush survives flooding and soil saturation longer than low sagebrush, and both silver Community composition reflects quality of and low sagebrush survive soil saturation forage, and community structure reflects better than big sagebrush. Tolerances for quality of cover. Foraging animals preferen- given environmental conditions vary even tially browse some sagebrush species and sub- within species: Bolander silver sagebrush is species more than others (Sheehy 1975; Smith more tolerant of alkalinity than is mountain 1950, 1952);cattle, domestic sheep, and prong- silver sagebrush and occurs in closed basins; horns favor black sagebrush more than big Wyoming big sagebrush is more tolerant of sagebrush (Dayton 1931). In another study, finer textured, alkaline soils than is basin big mule deer and domestic sheep preferred moun- sagebrush and therefore dominates it on such tain big sagebrush, Bolander silver sagebrush, sites. and low sagebrush; the mule deer ate but did not prefer Wyoming and basin big sagebrush The effect of climate on broad patterns of and showed the least preference for black sage- plant distribution is well accepted, and this brush (Sheehy 1975). Chemical composition effect is usually described and predicted by may influence the palatability of sagebrushes single factor relationships. For example, (Short et al. 1972). Sage grouse, horned larks, “Mean annual precipitation is never more than and pronghorns prefer low sagebrush stands

30 over big sagebrush stands +of the high desert butions are generally correlated with soil depth steppe. The taller big sagebrushes provide con- and seasonal wetness. Franklin and Dyrness siderable shelter from the elements for cattle, (1973). compiled information published by domestic sheep, mule deer, and sage grouse several authors and compared soils of four and nesting sites for many animals. Few mam- major sagebrush species: big sagebrush on mals and birds are small enough to effectively deeper soils, low sagebrush on shallow stony use the short sagebrush species for cover- soils, stiff sagebrush on very shallow stony thermal or hiding. Since many plant species of soils (lithosols), and silver sagebrush in the shrub-steppe often comprise both forage moister habitats. and cover, composition of plant species as well as crown cover of stands are measures of site Sagebrush communities of the Great Basin are valuable range habitats that help protect dominance and forage production. Height of as well as produce multiple resources. Sage- vegetation cover and depth of crown, and brushes comprise a protective cover that layering (adventitious rooting from branches minimizes wind and water erosion on many in contact with the soil) are attributes that sites. Grazing of livestock is the primary directly create thermal and hiding cover. measured economic use of the Great Basin Useful information is thus available from sagebrush zone. Free-ranging big game and identification of the community; its identifica- other wildlife provide many secondary benefits tion is accomplished by means of characteristic to society as well as an unmeasured but real plant species. This information can be applied economic flow within the region. Because of in evaluation, selection, and application of pro- these values, results from manipulating gram alternatives for management of that sagebrush vegetation are important to the stand (Daubenmire 1968, Kuchler 1957). short-term and the long-range economic goals of Western States. Every management strat- SagebrusWbunchgrass communities are egy in manipulating cover and forage qualities broadly distributed as a shrub-steppe type of the sagebrush range involves trade-offs in from Canada south through 11 Western States benefits to livestock and wildlife. into Mexico (Beetle 1960, Garrison et al. 1977). Extensive areas of the type are found in Washington (5.9 million ha or 14.7 million 11-BASIN BIG SAGEBRUSH/ acres), Oregon (10.9 million ha or 26.9 million BUNCHGRASS (fig. 22) acres), Idaho (15.0 million ha or 37.1 million Beetle (1960) described the range of basin acres), Nevada (13.7 million ha or 33.9 million big sagebrush in Oregon as extending from the acres), and California (7.3 million ha or 18.1 southern end of the Blue Mountains, north- million acres) (Beetle 1960). eastern Oregon, throughout the central and Stands vary from austere expanses domin- southeastern portions of the State. Although ated by single species to multihued mosaics he also recorded the basin form for every rich in species. county east of the Cascade Range, Winward (1980) considered its range more restricted Sagebrushes dominate arid, cold desert than that of other big sagebrush forms. The plains beginning at 30-meter (100-ft) elevation, land area occupied by basin big sagebrush con- but similar stands also mingle with moist, sub- stitutes a minor portion of the sagebrush com- alpine parklands up to 3 050 meters (10,000 ft) plex in Oregon; much of its former range is now in elevation. The stands are distributed down cultivated land. Over recent history, the distri- drainages, within basins, up slopes, across bution has changed little, but various uses plateaus, and along ridge crests. have considerably altered the density of stands Associated soils usually fall within either (Winward and Tisdale 1977). the Brown or Chestnut Great Soil Groups. This tallest of big sagebrush forms is Some sagebrush stands also occur on Cher- usually upright with erect spreading branches nozem or Sierozem soils (Fosberg and Hiro- and a definite trunk (fig. 21); occasionally it naka 1964, Tisdale et al. 1969). Species distri- may be either dwarf or treelike (Beetle 1960,

21 (1971), Dean (1960), Eckert (1957, 1958), Hansen (1956), Segura-Bustamante (1970), Urness (1966),and Volland (1976). Structure varies considerably among big sagebrush stands, and that variability influences their cover and forage qualities. Shrub height, crown cover, and plant density are commonly reported measures of structure of big sagebrush stands (table 7). Other important measures of structure seldom reported are the number and height of layers (vegetation strata) as well as hiding cover quality and crown depth. Many references contain structural measures, but most of these are concerned with big sagebrush in general-not a specific form, Figure 22.-A basin big sagebrushlbunch- such as basin big sagebrush (table 7). grass stand illustrating the typical structure of this sagebrush subspecies and an Site understory that has been intensively grazed for a long time, resulting in reduced In Oregon, basin big sagebrush is found density of bunchgrass and an increase of primarily along valley bottoms and in lower annuals. foothill regions between 30 and 2 140 meters (100 and 7,000 ft) in elevation. This subspecies Winward and Tisdale 1977). The flower stalks is also common to many sites with dry, shallow are scattered throughout the irregular top. The soils, southerly to westerly aspects, and at persistent leaves are long and relatively talus perimeters (Beetle 1960, Tisdale et al. narrow, shallow lobed, straight margined, and 1965, Winward 1980). Basin big sagebrush is widest at the lobe tips. Alcohol extracts of the also found in valley bottoms and lower foot- leaves fluoresce brownish-red in’ long-wave hills from 700 to 2 140 meters (2,500 to 7,000 ultraviolet light (Winward 1980). ft)in Idaho (Winward and Tisdale 1977). Vegetation Soi I Diversity of species in some stands creates Soil descriptions for many big sagebrush a rich composition, whereas other stands stands are readily available, but none were appear to be almost monotypes. Associated found that specified the subspecies of big plants that are common enough to be used in sagebrush that was present. Winward and naming big sagebrush communities and that Tisdale (1977), however, describe basin big also occur with basin big sagebrush are: Sand- sagebrush in Idaho as “growing in deep, well- berg bluegrass, needleandthread, Idaho fescue, drained soils. ’’ Our observations indicate the bearded bluebunch wheatgrass, giant wildrye, tallest stands of basin big sagebrush (to over and Thurber needlegrass (Daubenmire 1970, 2.4 m or 8 ft in height) grow in deep, well- Tisdale et al. 1965, Winward 1980). Franklin drained soils alongside rivers and streams in and Dyrness (1973, p. 235) compiled informa- southeastern Oregon. Big sagebrush/bunch- tion from several workers and demonstrated grass communities are common on shallow, the variety of plant species associated with big moderate, and deep soils. Textures include sagebrushes as well as the wide distribution of plants that identify the named communities silty clay loams through fine sandy loam and (table 6). Detailed species lists for big loamy sands to well-drained pumice sands sagebrushibunchgrass communities where the (Culver 1964, Daubenmire 1970, Dealy 1971, big sagebrush form is unspecified are available Fosberg and Hironaka 1964, Hall 1973, Tis- in Concannon (1978), Culver (1964), Dealy dale et al. 1965, Urness 1966, Volland 1976). The soil profiles generally show well-developed

32 Table 6-Characteristic species for Artemisia associations'

Species Association group High Lava Plains Owyhee Upland Southern Blue Mountains Artemisia tridentatd Shrubs Artemisia tridentata Artemisia tridentata Agropyron spicatum Grasses Agropyron spicatum Agropyron spicatum Poa sandbergii Poa sandbergii Herbs Phlox diffusa Lupinus sericeus Aster scopulorum Lomatium tritenatum Aster canescens Lomatium macrocarpum Chaenactis douglasii Zigadenus paniculatus Collinsia pamiflom Microseris troximoides Phlox gmcilis Astragalus filipes Lappula redowskii Astragalus lentiginosus Gayophytum ramosissimum

Artemisia tridentatd Shrubs Artemisia tridentata Artemisia tridentata Festuca iduhoensis Chrysothamnus viscidiflorus Chrysothamnus viscidiflorus Symphoricarpos rotundifolius Ribes cereum Juniperus occidentalis Grasses Festuca idahoensis Festuca idahoensis Agropyron spicatum Agropyron spicatum Poa sandbergii Poa sandbergii Koeleria cristata Sitanwn hystrix Bromus tectorum Elymus cinereus Herbs Phlox diffusa Balsamorhua sagittata Antennaria corymbosa Calochortus nitidus Artemisia tridentatd Shrubs Artemisia tridentata Elymus cinereus Grasses (mentioned but not described) Elymus cinereus Poa sandbergii Agropyron spicatum Bromus tectorum Herbs Penstemon speciosus Penstemon cusickii Thlaspi amense Eriogonum umbehtum

Artemisia arbusculal Shrubs Artemisia arbuscula Artemisia arbuscula Artemisia arbuscula Agropyron spicatum Eriogonum sphaerocephalum Purshia tridentata Juniperus occidentalis Grasses Agropyron spicatum Agropyron spicatum Agropyron spicatum Poa sandbergii Poa sandbergii Poa sandbergii Festuca idahoensis Sitanwn hystrix Sitanwn hystrix Bromus tectorum Herbs Phlox diffusa Penstemon aridus mfolium macrocephahm Erigeron linearis hgophyhmmosissima Collinsia parviflom

Artemisia arbusculal Shrubs Artemisia arbuscula Artemisia arbuscula Artemisia arbuscula Festuca idahoensis Juniperus occidentalis Grasses Festuca idahoensis Festuca idahoensis Festuca idahoensis Agropyron spicatum Agropyron spicatum Agropyron spicatum Poa sandbergii Poa sandbergii Poa sandbergii

Herbs Phlox diffusa Ambis holboellii Phlox douglasii Phlox hoodii Phlox diffusa Balsamorhua serrata Phlox longifolia Erigeron linearis Microseris troximoides Astragalus miser Antennaria dimorpha Balsamorhiza hookeri Astragalus stenophyllus Agoseris heterophyh Lupinus saxosus Achillea millefolium Trifolium gymnocarpon Haplopappus stenophyllus Trifolium macrocephulum

Artemisia rigidd Shrubs Artemisia rigida Artemisia rigida Poa sandbergii Grasses Poa sandbergii Poa sandbergii Bromus tectorum Sitanion hystrix Fes tuca micros tachys Agropyron spicatum Sitanwn hystrix Herbs Mimulus nanus Phlox douglasii Zigadenus paniculatus

'Adapted from Franklin and Dyrness (1973,p. 235). 33 Shrub height Ck0- Density Reference cover' Plants kea Square Square Meters Feet Percent Number meters feet

Big 0.4-2.0 1.2-6.6 - - - Hitchcock et al. (1955) sagebrush 1.1 3.5 13 3-7 9.29 Tisdale et al. (1965) 1.o -2.0 3.3-6.6 13-75 33 1076.00 Daubenmire (1970) .7 2.2 20-27 - - Culver (1964) .5-.6 1.7-1.9 7-12 11-15 18.58 Eckert (1957) .3-.7 1-2 7-27 - - Segura-Bustamante (1970) .4-.5 1.2-1.7 16-29 103-147 74.00 Urness (1966) 2-1.o .6-3.3 9.8-16.6 .3-4.7 10.76 Kornet (1978) - - 15-16 - - Volland (1976) - - 4-1 5 - - Hall (1973) Basin big .3-2.4 1-8 - - - Brunner (1972) sagebrush 1.0-5.0 3.3-16.0 - - - Beetle (1960) - - 13.2 ,057 .09 Sheehy (1975) 1.2-2.4 3.9-7.9 - - - Winward and Tisdale (1977)

Mountain big - - 12 .088 .09 Sheehy (1975) sagebrush .3-1.2 1.O -4.0 - - - Brunner (1972) 1.o 3.3 - - - Beetle (1960)

Wyoming big .3-.9 1.O -3.0 - - - Brunner (1972) sagebrush - - 8-23 - - Winward (1980) - - 9.7 .078 .09 Sheehy (1975) .4-1.0 1.5-3.3 - - - Winward and Tisdale (1977) Threetip .5 1.7 10 14 9.29 Tisdale et al. (1965) sagebrush 1.0-2.0 3.3-6.6 - - - Beetle (1960) 2-.6 .6-2.0 - - - Hitchcock et al. (1955)

Silver 1.5 4.9 - - - Beetle (1960) sagebrush .5-1.2 1.6-3.9 - - - Peck(1961)

Bolander .3-0.6 1.O -3.0 - - - Beetle (1960) silver - - 27.8 .33 .09 Sheehy (1975) sagebrush

Mountain 1.o 3.3 - - - Beetle (1960) silver sagebrush

~~ ~~ ~ ____ 'Same as crown canopy, canopy closure, or crown density-the proportion of ground surface covered by shrub crowns as vertically projected, like a shadow. horizons and often include a very fine-textured mule deer and domestic sheep preferred other B horizon. Brown and Chestnut soils are most subspecies to basin big sagebrush (Sheehy commonly associated with big sagebrush com- 1975, Winward 1980);basin big sagebrush was munities, but some stands have been found on never grazed in Nevada (Brunner 1972). other soils. Recognition of basin big sagebrush stands Discussion can aid the range manager in planning for maintenance or enhancement of cover and The forage value of big sagebrush has been forage. Crown cover may increase dramatically related to the quantity and quality of other because of crown enlargement after disturb- browse species within the same or adjacent ance, and in such communities there is a stands (Dietz and Yeager 1959; Short et al. greater potential for herbaceous production, 1972; Smith 1950, 1952).Use by wildlife varies native as well as introduced, than in some among taxa of big sagebrush. For example, other big sagebrush communities (Winward 1980).

34 Burning of various big sagebrushhmnch- sirable species, such as rabbitbrush, horse- grass stands produces different responses brush, goldenweed, and bottlebrush squirrel- among plant communities; different responses tail; burning produced grasslands that were among basin big sagebrush communities slowly reinvaded by big sagebrush (Volland should also be expected. Perennial grass cover 1976). is increased by burning of big sagebrush/ Thorough identification of a plant com- bearded bluebunch wheatgrass stands, but munity depends on the identification of its fescue plants are damaged by fire (Daubenmire species-an important step in determining its 1970, Concannon 1978). Fires eliminate big potential for management toward various sagebrush and initiate plant successions goals. This is often difficult to accomplish in during which perennial grass dominates the one visit to a stand because of differences in sites for long periods. timing of plant development among species. The influence of grazing in big Dean (1960) presented an account of peak sagebrushhnchgrass stands may vary rela- flowering times for species within big sage- tive to the dominant form of big sagebrush. brush types that illustrates this point. Daubenmire (1970) suggested that a big sagebrushlsandberg bluegrass community did 12,13-MOUNTAlN BIG SAGEBRUSH/ not result from overgrazing or burning of big BUNCHGRASS (fig. 23) AND SUBALPINE sagebrushlbearded bluebunch wheatgrass BIG SAGEBRUSHIBUNCHGRASS stands. In Oregon, the quantity of Sandberg The mountain big sagebrush subspecies is bluegrass increases as Idaho fescue declines, common in uplands from Oregon to the Rocky and this relationship is suggested as a measure Mountains (Beetle 1960, Winward and Tisdale of range condition (Tueller 1962). Sandberg 1977). At higher elevations, mountain big bluegrass, along with bottlebrush squirreltail sagebrush is replaced by subalpine big and longleaf phlox, increases as bearded blue- sagebrush, and an unnamed variant of moun- bunch wheatgrass declines in burned stands of tain big sagebrush extends from the lower edge the big sagebrushlbearded bluebunch wheat- of the western juniper zone into the steppe grass community (Concannon 1978). Tisdale et communities in northeastern and central al. (1969) found that big SagebrushlThurber Oregon (Winward 1980). Mountain big sage- needlegrass stands changed to big sagebrush is found in all Oregon counties east of brush/Sandberg bluegrass stands during the Cascade Range, from below Anthony heavy grazing. Soils associated with big sagebrush communities influence the use of stands by burrowing mammals, the rooting depth of plants, the patterns of soil moisture through the seasons, and the responses of plants after disturbance. Stone-free soils of big sagebrush/fescue stands were favorable to voles, ground squirrels, and badgers (Daubenmire 1970). Rooting depths were greater or effective moisture was better in big sagebrus hlgiant wildrye stands compared with adjacent stands of other communities (Culver 1964). Cooler and moister sites were indicated by big sagebrushlfescue stands, but soil moisture was depleted earlier in the needlegrass phase of the big sagebrushlbearded blue- Figure 23.-Mountain big sagebrushlbunch- bunch wheatgrass community (Eckert 1957). grass communities are species rich, have On pumice soils, manipulation of big sage- high forage production, and provide dense brush and other disturbances increased unde- cover for many wildlife species.

35 Lakes in Baker County southwest to the (1977) found the mountain form of big sage- Siskiyou Mountains in Josephine County brush on sites where annual precipitation (Beetle 1960, Winward 1980). varied from 0.5 meter (20 inches) to 1.1 meter (43 inches), slope varied from 6 to 33 percent, Mountain and subalpine subspecies of big and aspects included east through northeast to sagebrush are moderately tall, flat-topped shrubs (fig. 21B); the flower stalks come from north. Where both occur in high elevation only the upper part of the branches and extend areas, mountain big sagebrush is found on above the foliage (Beetle 1960, Winward and drier sites, subalpine big sagebrush on moister Tisdale 1977). The persistent leaves are wider sites (Winward 1940). relative to their length compared with basin big sagebrush, and they are broadest below the Soil leaf lobes (Winward and Tisdale 1977).Alcohol Mountain big sagebrush occurs on deep, extracts of the leaves of both the mountain and well-drained soils where moisture is available subalpine taxa fluoresce creamish-blue under throughout most of the summer (Winward and long-wave ultraviolet light (Winward 1980). Tisdale 1977). Schumaker and Hanson (1977) found this subspecies on mixed coarse and fine VggetaPion loamy soils from rhyolite and basalt as well as granite . Mountain big sagebrush/bunchgrass stands have a rich diversity of species; 40 plant associates are common (Winward 1980; also Discussion see footnote 7, p. 25). Some of the usual asso- The greater preference of ruminants for ciates are the same as those found with other mountain big sagebrush forage may be related big sagebrush taxa (table 6). Bearded blue- to phenological development, nutrient content, bunch wheatgrass, Idaho fescue, needleand- or essential oils. Mule deer have shown equal thread, giant wildrye, bottlebrush squirreltail, preference for mountain big sagebrush and its and Sandberg bluegrass are usual dominants unnamed variant,8 both being more preferred in the grass-forb layer. Prairie junegrass and than basin or subalpine big ~agebrush.~Sheehy Kentucky bluegrass are more usual with the (1975) found that mule deer and domestic subalpine form than with mountain big sage- sheep preferred Wyoming big sagebrush over brush. Rabbitbrushes are usually present, and mountain and basin big sagebrush. Brunner their densities are greater in disturbed stands. (1972) found that mountain big sagebrush was California brome, slender wheatgrass, onion- seldom grazed in Nevada. Growth of mountain grass, and sedges are common associates of big sagebrush is initiated later than that of subalpine big sagebrush (Winward 1980). basin and Wyoming big sagebrushes by about Mountain big sagebrush and the subalpine 2 weeks, but seed ripening of the mountain form have structurally dense stands (table 7); subspecies precedes that of the other two they have the potential for attaining greater subspecies by about 2 weeks (Winward and crown cover than the other sagebrush species, Tisdale 1977). Cattle and domestic sheep subspecies, or form. Crown cover increases grazed sagebrush principally in spring and fall after disturbance and as range condition according to Garrison et al. (1977), but deteriorates (Winward 1980, Winward and Pechanec et al. (1965) found that sheep also Tisdale 1977). depended on sagebrush for winter browse. Mountain big sagebrush shows the least variation in chemical content and also has the most Sits persistent leaves among sagebrush taxa Many upper foothill and mountain sites (Beetle 1960). The chemical value of different from 1 070 to 3 050 meters (3,500to 10,000 ft) 8Persona1 communication from A. H. Winward, Oregon in elevation are occupied by mountain big sage- State University, Corvallis. brush (Beetle 1960, Winward 1980, Winward ePersonal communication from D. P. Sheehy, Oregon and Tisdale 1977). Schumaker and Wanson Department of Fish and Wildlife, Burns.

36 sagebrushes compares favorably with other forages, although pure diets of sagebrush appear inadequate for maintenance of animals (Beetle 1960, Cook 1972, Dietz and Yeager 1959, Dietz et al. 1962). Consumption of sagebrush foliage may be limited by antimicrobial substances, such as essential oils, but in- hibitory effects on the rumen microflora demonstrated in the laboratory (Nagy and Ten- gerdy 1968) are not likely to be attained in nature because of insufficient rates of release in the acid environment of the rumen (Short et al. 1972). Perhaps the varying consumption of sagebrush forms by grazers, however, is related to differences in consumption and content of essential oils among the sagebrushes Figure 24.-Wyoming big sagebrush/bunch- (Sheehy 1975). grass communities occur on relatively dry The dense structure of mountain and sites and are the most common of the subalpine big sagebrush stands and their sagebrush-dominated communities in the capacity for increases in plant density and Great Basin of southeastern Oregon (Chris Maser photograph). crown cover after disturbance make these communities valuable cover habitats for wildlife. shrub, 0.4 to 1 meter (1.5 to 3.3 ft) tall, with an Managers should plan periodic control of parts irregular and rounded top (fig. 21A) and with of some stands to develop openings for mule the flower stalks scattered throughout the deer, thus producing more diversity and crown-like small shrubs of basin big sage- greater edge length. In fawning and fawn- brush. The main stem is not trunklike, how- rearing areas dominated by mountain big sage- ever, but is comprised of two or three twisted brush or subalpine big sagebrush, some dense portions at ground level. The persistent leaves stands of either form may be best left alone are relatively short and wide, more deeply (Winward 1980). The mountain subspecies is lobed. than in other big sagebrush taxa, and readily controlled by flooding (Beetle 1960), as their margins curve outward from the base to are other forms of big sagebrush (see footnote form bell-shaped blades. Alcohol extracts from 8). the leaves fluoresce shades of brownish-red in Seeding managed stands of mountain big longwave ultraviolet light (Winward 1980, sagebrush with crested or intermediate wheat- Winward and Tisdale 1977). grass and seeding stands of subalpine big sagebrush with intermediate wheatgrass, Cali- Vegetation fornia brome, or smooth brome are usually suc- Many plant species commonly occurring cessful. Seeding, however, is infrequently with Wyoming big sagebrush are also found required becausel the understory is usually in with other taxa, such as basin or mountain big good condition (Winward 1980). Large popula- sagebrush. Major associated species include: tions of goph.ers in some dense stands of sub- bearded bluebunch wheatgrass, needleand- alpine big sagebrush sufficiently disturb the thread, Thurber needlegrass, bottlebrush soil to increase the annual forb component. squirreltail, Sandberg bluegrass, and cheatgrass. Idaho fescue occurs occasionally with 14-WYOMING BIG SAGEBRUSH/ Wyoming big sagebrush (Schumaker and Han- BUNCHGRASS (fig. 24) son 1977, Winward 1980). Many stands have a Wyoming big sagebrush is the most sparse grass-forb layer because of heavy use by common sagebrush throughout the high desert livestock and wildlife; some stands and the in Oregon (Winward 1980). It is a low growing understory have been altered by natural,

37 periodic burning (Winward 1980). These distur- palatable as antelope bitterbrush and often bances and loss of understory are associated severely grazed in parts of Nevada (Brunner with increased density of sagebrush or other 1972). Sparse grass-forb layers are common in shrubs. There are few perennial forbs, antelope Wyoming big sagebrus Wbunchgrass stands, bitterbrush is not a natural component, and and they offer little forage from associated cryptogams may fill much of the bare areas in plant species. undisturbed stands. The lateral rooting of this subspecies may compete more with herbaceous Dense, low cover for small animals is of- species than that of other big sagebrush taxa fered by some stands of Wyoming big (Winward and Tisdale 1977). A correlation sagebrush. Often, however, the shrubs are too between crown cover or crown diameter and small or scattered to provide much protection, production has been determined for this sub- especially not for large mammals and birds. species (Rittenhouse and Sneva 1977). The Disturbances of the grass-forb layer causes on- stands are not structurally dense (table 7), yet ly moderate increases in density of this they may totally occupy the site. subspecies.

Site 15-THREETIP SAGEBRUSH/ BUNCHGRASS (fig. 25) Wyoming big sagebrushlbunchgrass is Threetip sagebrush is distributed in Wash- most common at elevations of less than 1 830 ington, Idaho, and Oregon (Beetle 1960, meters (6,000 ft) and on more xeric mountain Daubenmire 1970). Within Oregon, the species sites than other big sagebrush communities occurs in Baker and northern Harney counties (Winward 1980). Schumaker and Hanson (1977)described stands on northeast aspects of and occasionally in Malheur County (Beetle 8-percent slope in an area with 33 centimeters 1960, Winward 1980; also see footnote 8, p. 36). (13 inches) of annual precipitation. The subspecies is common on slopes of major drainages in hot and dry areas from 700 to 1 980 meters (2,500 to 6,500 ft) in Idaho (Winward and Tisdale 1977). In Nevada, this subspecies is found within the terrace sagebrush complex (Brunner 1972).

Soil Relatively shallow to moderately deep soils are present under stands of Wyoming big sagebrush/bunchgrass; often the soil is slightly calcareous in the surface layer (Winward 1980, Winward and Tisdale 1977). Fine, loamy soils from basalt were associated with the com- Figure 25.-Threetip sagebrushlbunchgrass munity in Idaho (Schumaker and Hanson stand in a usual topographic location. The 1977); the community is also found on “red- elevation of this stand, 865 meters rock surfaces” over deep soils in Nevada (2,837 ft), in northern Malheur County, (Brunner 1972). Oregon, is lower than reported in the literature (see footnote 5, p. 18). Discussion Threetip sagebrush is an erect shrub (fig. Wyoming big sagebrush was of low to 21A) that freely and profusely branches to intermediate palatability for mule deer and form a moderately large crown, 0.4-0.8 meter domestic sheep in Oregon compared with six (1.3-2.6 ft) in diameter (Beetle 1960). This other sagebrush taxa (Sheehy 1975, Winward species does not normally root from its 1980). Conversely, this subspecies was as branches (layering)because of its upright form;

38 it readily does so when the branches touch the slopes and in depressions, whereas threetip soil, and it sprouts readily from burned crowns sagebrushlbearded bluebunch wheatgrass (Beetle 1960). Dwarf threetip sagebrush is a stands are found on dry, warm sites where ex- low subspecies in some stands, is rarely taller posure to sun and wind is too severe for fescue; than 0.2 meter (0.7 ft), has a crown from 0.3 to for example, on east slopes curving to the 0.4 meter (1.0 to 1.3 ft) wide, and readily layers north there are abrupt ecotones between the from its naturally decumbent branches (Beetle communities (Daubenmire 1970, Tisdale et al. 1960). This subspecies is usually found adja- 1965). In Oregon, this species is reported at 'cent to mountain big sagebrush but not elevations of 1 160 meters (3,800 ft)'or higher mingled with it; dwarf threetip sagebrush by Winward (1980). Dealy (see footnote 5, occupies shallow soils along ridges rather than p. 18) reports a low elevation of 865 meters the deeper soils at the base of foothills (Beetle (2,837 ft) for this species in northern Malheur 1960). County, Oregon. Beetle (1960) reported that threetip sagebrush occurred between 914 and The leaves of threetip sagebrush are rela- 1 830 meters (3,000and 6,000 ft). tively long, 3 centimeters (1.2 inches), and deeply three cleft into linear lobes which may Soi I also be divided (Beetle 1960). Alcohol extracts of the persistent leaves fluoresce creamish-blue Threetip sagebrush is the only resprouting under long-wave ultraviolet light (Winward sagebrush species associated with loam to 1980). sandy-loam soils in Oregon (Winward 1980). Daubenmire (1970) found stands of the three- Vegetation tip sagebrushlbunchgrass community on Brown, Chestnut, Chernozem, Planosol, and Many plant species found with big sage- Prairie soils in Washington and Idaho. Three- brushes in general, occur with threetip tip sagebrush communities were associated sagebrush in its fescue and wheatgrass com- with silty clay loams containing a B horizon at munities (Daubenmire 1970, Tisdale et al. 0.2-0.3 meter (0.7-0.8 ft) and a lime zone below 1965, Winward 1980). Plant species composi- 0.9 meter (2.9 ft) in Idaho (Tisdale et al. 1965). tion for the threetip sagebrushlfescue and In general, threetip sagebrush communities threetip sagebrushlwheatgrass communities are associated with deep, well-drained soils were similar except for the occurrence of (Beetle 1960). fescue. Extensive species included the following: rabbitbrush, big sagebrush, antelope bit- Discussion terbrush, squaw apple, Sandberg bluegrass, bearded bluebunch wheatgrass, prairie june- Apparently the intensity of browsing of grass, needleandthread, Thurber needlegrass, threetip sagebrush varies by subspecies and Kentucky bluegrass, threadleaf sedge, arrow- areas. Winward (1980)considered that grazing leaf balsamroot, dwarf hesperochiron, longleaf by native and domestic animals contributed to phlox, nineleaf biscuitroot, and tapertip the density of present threetip sagebrushl hawksbeard. bunchgrass stands. Brunner (1972) observed that threetip sagebrush was never grazed, but Canopy cover measurements are high he reported that dwarf threetip sagebrush was where plants are dense (table 7), since in- usually severely grazed. dividual crowns are of large diameter, 0.4 to 0.8 meter (1.3 to 2.6 ft), and freely branched The structure of threetip sagebrushlbunch- (fig. 21A). Threetip sagebrush may form grass stands should provide considerable stands sufficiently dense to crowd out her- cover, but the shorter dwarf threetip sage- baceous species (Winward 1980). brush can protect only smaller animals. The resprouting ability and growth form can pro- Site duce very dense brush fields that vigorously compete with herbaceous species (Winward Threetip sagebrush/fescue stands occur on 1980). Stable shrub densities cannot be main- moist and cool sites, such as north and east tained with proper grazing or even no use;

39 periodic thinnings are necessary to attain some management goals. Sandberg and Kentucky bluegrasses are common in threetip sagebrush/bunchgrass stands, and crested wheatgrass can be established on these sites (Winward 1980). The stone-free mound soils associated with some threetip sagebrush communities provide a favorable substrate for burrowing rodents and badgers (Daubenmire 1970).

16,17-BQLANDER SILVER SAGEBRUSH/ BUNCHGRASS (fig. 26) AND MOUNTAIN SILVER SAG EBRUSHlBU MCHG RASS Figure 26.-This stand of Bolander silwer (figs. 27 and 28) sagebrushlbunchgrass on an internally drained basin site illustrates the usual Silver sagebrushes are found in seasonally bare soil interspaces and sparse moist areas from central and eastern Oregon understory. eastward across the Rocky Mountains to Nebraska and the Dakotas. Two subspecies are associated with two distinct habitats, closed basins and streamside or pond-edge meadows, that are scattered throughout the Pacific Northwest and intermountain regions (Beetle 1960). The basin subspecies, Bolander silver sagebrush, is distributed within the desert areas of Oregon from Prineville throughout the southeastern third of the State; the streamside-meadow subspecies, mountain silver sagebrush, is most common in east-central and southeastern Oregon where there are sea- Figure 27.-Mountain silver sagebrush/ sonally high water tables adjacent to streams bunchgrass stand adjacent to a mountain and meadows (Dealy 1971, Winward 1980). meadow. These two silver sagebrush subspecies differ morphologically, chemically, and eco- logically. Bolander subspecies is a low rounded shrub with erect or spreading dense branches, but mountain silver sagebrush can be taller and more erect (Beetle 1960). Branching of both subspecies creates an irregular crown (fig. 21A). Branches of both subspecies readily form roots when they touch the soil, and the plants resprout from the root crown. The leaves are tapered toward both ends, are usually not lobed, are very silky with white hairs, and smell pungent (like turpentine) when crushed (Beetle 1960). Alcohol extracts of leaves from either subspecies fluoresce creamish-blue shades under long-wave ultraviolet light (Win- Figure 28.-This detailed wiew of a mountain si Iwer sagebrus hlbu nc hgrass stand i I I u- ward 1980). The unlobed leaves and tolerance strates the usually dense understory.

40 of soil saturation and seasonal flooding clearly 0.3-0.6 meter (1-2 ft) for at least a month each set both silver sagebrushes apart from all spring when the snowpack melts. There have other shrubby, tall sagebrushes. Leaves of the been years-1965, for example-when flooding other tall species are cleft three or more times, persisted even beyond the tolerance of silver and those shrubs are easily killed by even brief sagebrushes and aerial portions of the shrubs periods of flooding or soil saturation. died. The dead branches were replaced within 3 years by new or resprouted individuals, after Vegetation the basins passed through a structural condi- Moist sites dominated by .silver sage- tion dominated by Newberry cinquefoil and brushes support rich herbaceous understories annual forbs (see footnote 7, p. 25). Sites con- in spring, but these understories are usually taining the mountain subspecies usually are maintained through the grazing season only in saturated in spring and early summer, but nor- mountain silver sagebrush stands. The sage- mally they are covered only briefly with stand- brushes are usually the only shrubs present, ing water. The exception is found in wet but muhly, spike rush, rushes, sedges, and a meadows that form ponds (Dealy 1971). Both few forb species are associated (Dealy 1971, subspecies occur at midelevation to high elevation: Bolander silver sagebrush, 1 000-1 800 Winward 1980; also see footnote 7, p. 25). meters (3,500-6,000 ft); mountain silver sage- Other plants commonly observed in Bolander brush, 1 680-2 240 meters (5,500-8,000 ft) silver sagebrush stands include: Douglas (Beetle 1960, Winward 1980; also see footnote sedge, Baltic rush, Newberry cinquefoil, bottlebrush squirreltail, showy downingia, combleaf, 8, p. 36). tiny mousetail, and cheatgrass (Leckenby and Soil Toweill 1979, Sheehy 1975). Herbaceous species are more abundant in the floristically Silver sagebrush communities are asso- rich mountain silver sagebrush stands and in- ciated with deep soils of variable surface tex- clude some species not present in Bolander ture (sand to clay) (Beetle 1960, Cornelius and silver sagebrush communities, such as: slender Talbot 1955). Within Oregon, however, these wheatgrass, California brome, Nevada subspecies are dominant on deep clay and bluegrass, sedges, timothy, and Kentucky alkaline soils (Leckenby and Toweill 1979, bluegrasslo (Winward 1980). Urness 1966, Winward 1980). Clay soils found with mountain silver sagebrush develop wide Structural attributes of free branching, deep cracks as they dry over the summer, and layering, and crown sprouting can create con- the flaking surface layers readily slough into siderable cover, but silver sagebrushes are deep fissures (Cornelius and Talbot 1955; also seldom sufficiently tall (table 7) to provide see footnote 10). cover for animals larger than swans, geese, rabbits, and coyotes. Discussion Site Use of silver sagebruddbunchgrass stands varies among seasonal ranges. Bolander silver Subspecies of silver sagebrush are closely sagebrush foliage is highly preferred by mule associated with sites that are seasonally very deer and domestic sheep (Sheehy 1975; also see moist to saturated or even temporarily flooded footnote 7, p. 25). These basin communities (usually late winter and spring) (Beetle 1960, are occupied by mule deer more than expected Cornelius and Talbot 1955, Dealy 1971, Win- from their availability on the Silver Lake-Fort ward 1980; also see footnote 7, p. 25). This Rock winter range (Leckenby 1978a, Urness single environmental condition' eliminates 1966). On summer range in the same area, most other plants common to shrub-steppe Dealy (1971) considered silver sagebrush/ communities. The internally drained, alkaline muhly stands of low value because they con- basins dominated by Bolander silver sage- stituted a small area and thus provided limited brush are frequently ponded to depths of forage compared with that in adjacent com- loSchallig,W. H. C., unpublished data on file at Oregon munities. Dealy (1971), however, found that State University, Corvallis.

41 competition between livestock and deer ap- peared high for the limited herbaceous forage available. All silver sagebrush communities should be considered important foraging habitats for mule deer, pronghorns, and sage grouse. Depth and density of vegetation may provide good cover in some mountain silver sagebrush stands, but cover is sparse in the severely grazed alkaline basins dominated by Bolander silver sagebrush. Pronghorns frequently use these “flats” as resting and bed- ding areas throughout the summer. The heavy soils and seasonal ponding make Figure 29.-The site on which this sparse forage improvement in most silver sagebrush stand of stiff sagebrushlbunchgrass oc- sites risky at best; introduced species have curs illustrates the severe conditions rarely survived well (Cornelius and Talbot under which the species can grow in the 1955, Leckenby and Toweill 1979, Winward Great Basin of southeastern Oregon. 1980).These communities are best left untilled Oregon. It is common in the following Oregon and unsprayed because of their wildlife value counties: Wasco, Wheeler, Crook, Gilliam, Jef- and because the native plants are difficult to ferson, Umatilla, Union, Wallowa, and Harney replace with other forage species that do as (Beetle 1960). Winward (1980) found stiff well. sagebrush stands distributed primarily in Waterholes are often developed in silver northern and northeastern Oregon, but stiff sagebrush basins. Frequently, ditches are dug sagebrush was also located from northern across the basin floor to drain a shallow pond Harney County east through Malheur County rapidly into a deep reservoir which maintains a into Idaho. reliable water source longer through the live- Stiff sagebrush is a winter deciduous, low, stock grazing season. The improved drainage spreading, and dense shrub (fig. 21C) that con- produces a shorter period of flooding; new, sists of short, rigid, decumbent, and brittle drier, plant habitats are created that favor big branches (Hitchcock et al. 1955, Peck 1961, sagebrush and rabbitbrushes over silver Winward 1980). Alcohol extracts of the sagebrush. In smaller basins, silver sagebrush narrow-based and tridentlike lobed leaves stands are being slowly replaced by big fluoresce brownish-red under long-wave ultra- sagebrush-rabbitbrush communities (Leck- violet light (Winward 1980). enby 1978a). Thus, some trade-offs among multiple resource goals are illustrated in the Vegetation case of silver sagebrush basins altered for enhancement of water sources for livestock. Stiff sagebrushibunchgrass plant communities are often floristically rich but may appear impoverished because of spacing be- 18-STIFF SAGEBRUSHlBUNCHGRASS tween plants and sparse ground cover (fig. 29) (Daubenmire 1970). Common associated plant Stiff sagebrush (also called scabland sage- species include several mosses (TortuLa, brush) communities occur from the Cascade Bryum, C‘eratodon, Grimmia), woodlandstar, Range and Blue Mountains of Washington biscuitroot, spring draba, autumn willowweed, through central and southeast Oregon into pink microsteris, dwarf monkeyflower, Sand- Idaho (Daubenmire 1970). This short sage- berg bluegrass, cheatgrass, Pacific fescue, brush species is found regularly at the bearded bluebunch wheatgrass, and bottle- northern end of the Great Basin within eastern brush squirreltail (Culver 1964, Daubenmire

42 1970, Winward 1980). Hall (1973) found dwarf Soil squirreltail, false agoseris, a biscuitroot, and Stiff sagebrush communities are associated bighead clover common to stiff sagebrush with very shallow to shallow 10- to 28-cen- stands in the Blue Mountains. Other common timeter (4- to 11-inch), stony soils that have associated species are listed in table 6. been developed from basalt and rhyolite Although many species provide diversity (Culver 1964, Daubenmire 1970, Hall 1973, of forage, the small shrubs and open stands Winward 1980). Soil textures vary from loams provide little cover for other than the smallest to fine clay loams. The profiles usually become animals (table 8). saturated with water in winter and spring and are regularly subjected to frost heaving or Site frost boils. Stiff sagebrush in the Great Basin of Oregon is found exclusively on various aspects Discussion of rocky scablands that have undulating or Big game and livestock use stiff sagebrush rolling relief. Stiff sagebrushlbunchgrass as browse (Daubenmire 1970, Hall 1973, Win- stands are found from 914 to 2 134 meters ward 1980; also see footnote 7, p. 25). Many (3,000 to 7,000 ft) in elevation. They are commonly associated species are valuable usually on gentle slopes or benches of 0- to 20- forage for grazing animals. percent slope and occasionally on steeper slopes of up to 40 percent (Beetle 1960, Culver The low stature and dispersion of stiff 1964, Hall 1973, Winward 1980). sagebrush shrubs does not provide cover of any consequence for .animals larger than

Table 8-Structural measurements of short sagebrushes

Sagebrush Shrub height Crown Density Reference species ' cover' Plants Area Square Square Meters Feet Percent Number meters feet

Stiff 0.2-0.4 0.6-1.3 - Peck (1961) - - 20 Culver (1964) - - 10-34 Daubenmire (1970) - - 5-20 Hall (1973)

Low .l-.4 .3-1.3 13-16 Eckert (1957) - - 26 Culver (1964) .l-.2 .4-.5 4-22 Kornet (1978) - - 10.8 Sheehy (1975) - - 10 Volland (1976) .2-.3 .7-.9 6-20 Segura-Bustamante (1970) .2 .7 8-26 Dealy (1971)

Black - - 11.8 Sheehy (1975) .l-.3 .3-1.O - Beetle (1960) .l-.4 .3-1.3 - Hitchcock and Cronquist (1973) .3 1.o - Brunner (1972) .l-.3 .3-1.0 - Dayton (1931)

Early low .2 .8 1.5 Tisdale et al. (1965) .3 1.o - Beetle (1960)

'Same as crown canopy, canopy closure, or crown density-the proportion of ground surface covered by shrub crowns as vertically projected, like a shadow.

43 horned larks or ground squirrels. The lack of leaves (this is the only deciduous shrubby sagebrush in the area) in winter severely reduces the little cover this species offers on the scablands during periods of thermal stress. Even in winter, however, the shrubs do provide some protection from erosion by wind (Hall 1973). Stressful environmental conditions for plant growth, such as waterlogging and consistent frost heaving of very shallow soils, make successful seedings of domestic grasses highly improbable (Hall 1973, Winward 1980). Re- moval or control of the stiff sagebrush cover Figure 31.-This stand of cleftleaf would increase thermal stress for small sagebrushlbunchgrass occurs at 1 524 animals, reduce the forage available for both meters (5,000 ft) in east-central Malheur large and small animals, and increase erosion County, Oregon, adjacent to curlleaf by wind. mountainmahogany. This is a new location in Oregon for the subspecies (see footnote 19,20-L0W SAGEBRUSHlBUNCHGRASS 5, p. 18). (fig. 30) AND CLEFTLEAF SAGEBRUSH/ BUNCHGRASS (figs. 31 and 32) Low sagebrush/bunc hgrass communities typically occur adjacent to or intermixed with big sagebrush communities but are distinctly separate stands of an edaphic (soil-related) climax vegetation associated with shallow, stony soils (Dealy 1971, Dealy and Geist 1978, Franklin and Dyrness 1973). Low sagebrush is common in most counties east of the Cascade Range in Oregon. Beetle (1960)listed the counties where low sagebrush occurs (Baker,Grant, Crook, Jefferson, Wheeler, Harney, Malheur, Lake, Klamath, and Jackson. Figure 32.-Closeup of a cleftleaf sagebrushlbunchgrass stand illustrating the habit (general appearance) of this subspecies of sagebrush and the gravelly, shallow soil of sites on which it grows in the Great Basin of southeastern Oregon.

Low sagebrush is a gray to green dwarf shrub formed of irregular, short, and stiff branches (fig. 21C). It produces a small crown between 0.4 and 0.8 meter (1.3 and 2.6 ft) wide (Beetle 1960, Brunner 1972). The species infrequently forms roots from branches touching the soil. Both the small, narrow, wedge-shaped, Figure 30.- Low sagebrushlbunchgrass communities are common on shallow, and deeply three-toothed leaves of low sage- stony soils (photo, courtesy Oregon brush, as well as the much thinner lobed leaves Department of Fish and Wildlife). of the subspecies cleftleaf sagebrush, fluoresce

44 creamish-blue in alcohol solutions exposed to uplands, top-of-rim edges, level and sloping long-wave ultraviolet light (Winward 1980). A plateaus, and crests and slopes of ridges taller variant (as yet unnamed) associated with (Culver 1964; Dealy 1971; Dean 1960; Eckert pumice soils requires more research (Winward 1957, 1958; Hall 1973; Segura-Bustamante 1980.) 1970; Volland 1976; Winward 1980).

Vegetation Soil Stands vary from small-2 hectares (5 Low sagebrush thrives on shallow, stony, acres)- to wide “flats”l.6 kilometers (1mi) or fine-textured soils derived from basaltic, more across. The usual associated plants andesitic, or rhyolitic parent materials. These create a rich diversity of species within these soils may have basic, neutral, or acidic reac- stands of vegetation. Common associates are: tions. They generally are less than 0.6 meter (2 bearded bluebunch wheatgrass, Idaho fescue, ft) deep, may contain an impermeable (or at , Thurber needlegrass, Sandberg bluegrass, least restrictive) clay B horizon, become prairie junegrass, one-spike oatgrass, bottle- saturated with water in late winter and spring, brush squirreltail, cheatgrass, western needle- and are extremely droughty in summer (Brun- grass, woolly eriophyllum, Bloomer fleabane, ner 1972, Culver 1964, Dealy 1971, Dealy and low pussytoes, yarrow, gay penstemon, Geist 1978, Eckert 1957, Fosberg and Nevada biscuitroot, Holboell rockcress, Hironaka 1964, Hall 1973, Segura-Bustamante starved milkvetch, obscure milkvetch, spread- 1970, Volland 1976, Winward 1980; also see ing phlox, longleaf phlox, Hooker balsamroot, footnote 7, p. 25). Lack of physical support annual agoseris, daggerpod, granite gilia, during spring periods of soil saturation can bighead clover, and nineleaf biscuitroot result in damage from trampling (Hall 1973). (Culver 1964; Dealy 1971; Dean 1960; Eckert Extremes of saturation, frost heaving, and 1957, 1958; Hall 1973; Segura-Bustamante drying in these soils make plant survival 1970; Volland 1976; Winward 1980; also see tenuous at best. Though the species are well footnote 7, p. 25). adapted, even the low sagebrush and Sandberg As with other structurally short sage- bluegrass roots are pedestaled and broken dur- brushes, low sagebrush shrubs are too small ing seasonal cycles of frost heaving and soil and too scattered to provide cover for large drying. Often, cracks in the underlying base animals (table 8), but the dense crowns do rock, or interrupted restrictive layers in the shelter small animals, such as lizards, snakes, solum produce dispersed soil microsites that birds, and mice. The evergreen low sagebrush are deeper and better drained than surround- does, however, maintain its minimal cover ing soil. This permits establishment and sur- qualities through winter better than does the vival of a few scattered ponderosa pine, deciduous stiff sagebrush. western juniper, curlleaf mountainmahogany, antelope bitterbrush, or other shrubs (Dealy Site and Geist 1978, Segura-Bustamante 1970). Low sagebrushlbunchgrass communities Discussion are regularly found on dry, relatively sterile, often alkaline sites (Beetle 1960). Although Low sagebrush stands are intensively used Beetle (1960) reported low sagebrush often by wildlife and are particularly important to occurring on alkaline sites, Brunner (1972),pre - large ruminants, including livestock. Mule senting data on the sagebrush genus in deer regularly and preferentially occupied Nevada, and Dealy (1971), studying low sage- these communities during mild weather in brush in Oregon, report low sagebrush only on winter and spring (Leckenby 1978a). Use by pronghorn and sage grouse is also high, though acid to neutral sites. Occurrences are between 914- and 2 743-meter (3,000- and 9,000-ft) less seasonal. Indigenous species of wildlife elevation. The stands are on most aspects, but intensively graze the associated forbs and commonly they are found on gentle slopes (2-15 grasses. Browsing of low sagebrush by insects, percent) in rolling to undulating or flat mice, rabbits, hares, sage grause, and ruminants is also extensive. \

45 Forage species develop as much as 2 weeks compared with adjacent big sagebrush com- earlier in low sagebrush stands than in the munities of similar composition (Segura- adjacent antelope bitterbrush, tall sagebrush, Bustamante 1970): western juniperlantelope rabbitbrush, western juniper, curlleaf moun- bitterbrush -big sagebrush, 39-percent total tainmahogany, or ponderosa pine communi- cover; western juniperlantelope bitterbrush- ties. Grazing animals follow the sequence of low sagebrush, 38 percent; antelope bitter- forage development which is induced by dif- brush-big sagebrush, 36 percent; antelope ferences in site factors among these communi- bitterbrush-low sagebrush, 34 percent; big ties. Similar grazing patterns exist elsewhere sagebrush, 35 percent; and low sagebrush, 23 and appear to be an expression of resource par- percent. Crown cover of western juniper, titioning related to rates of plant growth which however, was similar among stands dominated in turn are correlated with different plant by either sagebrush. Abundance of forbs in- habitats. In a shortgrass zone, deBoer (1974) creased steadily, whereas total crown cover found that herds of grazing wildlife preferred decreased in those stands. Densities of bearded plant communities produced by a shallow soil bluebunch wheatgrass, Thurber needlegrass, overlying a restrictive hardpan (a soil environ- Sandberg bluegrass, and Idaho fescue were ment similar to that of short sagebrush greater where these grasses occurred with low stands). Low total herbaceous production and sagebrush compared with big sagebrush/ low rates of growth on these sites apparently bunchgrass stands (Segura-Bustamante 1970). permitted earlier season grazing that was Management designed to harvest or im- nonselective for plant species and that main- prove production of forage from low sage- tained growth of vegetative parts that were brush/bunchgrass stands should be planned highly digestible. Two other vegetational after careful evaluation of the goals, trade-offs, zones, both characterized by more precipita- and risks. These sites are fragile, generally will tion, deeper soils, and lack of a restrictive not produce much more forage after treatment, layer, produced higher rates of plant growth are not suitable for cultivation, and occur on (soil environments similar to those of tall soils that are too shallow for crested sagebrushes and other steppe communities). wheatgrass or other readily available intro- There, the grazing animals exhibited marked duced species (Dealy 1971, Hall 1973, Volland selectivity of forage species and grazed these 1976, Winward 1980). Abundance of remnant stands much later and less intensively than forbs and grasses may improve with changes those of the shortgrass zone. in grazing management (Dealy 1971, Winward Low sagebrush was one of the most pre- 1980), but some stands in poor condition have ferred sagebrushes offered to mule deer and apparently not changed even where they were domestic sheep (Sheehy 1975). Some sub- completely protected for 30 years or more. species of low sagebrush are grazed more Identification of specific low sagebrush extensively than others by mule deer and other plant communities requires data from more wildlife (Brunner 1972, Dealy 1971, Leckenby than one season (Leckenby 1978a).Dean (1960) 1978a, Volland 1976, Winward 1980). In addi- described plant species appearance (especially tion, the plant composition of many stands phenology of flowering) in spring, summer, and offers a rich diversity of seasonal forages. autumn. The marked differences in time of Height, crown cover, and plant density of plant species appearance, recognition, or prom- low sagebrush/bunchgrass stands provide inence in the physiognomy (outward feature) of little structure to create hiding or thermal a stand produce quite different impressions of cover for animals larger than ground squirrels, low sagebrush communities. In spring when mice, and small birds; these communities are obscure milkvetch is prominently flowering, primarily habitats for production of forage. the foliage easily contrasts with the wet soils; Total crown cover of all vegetation was less in summer when the colorful ashy penstemon where low sagebrush/bunchgrass stands were is in full bloom, the obscure milkvetch has all but disappeared.

46 21 -BLACK SAGEBRUSHlBUNCHGRASS 1980). The species is common to level plains and slopes in foothills and mountains and is Black sagebrush is more common in that reported to occur with ponderosa pine, aspen, portion of the Great Basin east of Oregon and woodland types-even into the spruce-fir (Hitchcock et al. 1955), but Beetle (1960) reported it scattered in Oregon’s Harney and zone (Dayton 1931). Lake Counties. Winward (1980)found that the relatively few areas of this sagebrush occurred Soil primarily in the southern portions of Malheur, The black sagebrushlbunchgrass com- Harney, and Lake Counties. West et al. 41978) munity, like other short sagebrushes, is observed black sagebrush in many stands associated with shallow, stony soils (Beetle throughout Nevada and in Utah. 1960). This species, however, has particular The shrub is low to dwarf (fig. 21C); its affinity for calcareous soils (Brunner 1972, branches are decumbent; the flower stalks are Winward 1980). Either lime hardpans or a con- brown and persistent (Beetle 1960). Although centration of lime is present in most soils some authors consider black sagebrush a supporting this sagebrush species. Such a relationship contrasts with that of the neutral to variety of low sagebrush (Hitchcock and Cron- acid volcanic soils common to low sagebrush quist 1973), there are important differences stands (Brunner 1972, Dealy 1971). (Winward 1980). There may be subspecies of black sagebrush that are also important to Discussion recognize, for at least a glossy-green form appears to be grazed less than a gray-green Six other sagebrush species were more form (Brunner 1972). Beetle (1960) also palatable than was black sagebrush to mule described these color phases. The broadly deer and domestic sheep (Sheehy 1975). Prong- wedge-shaped (flabelliform), three- to five- horns (Winward 1980), domestic sheep, and lobed leaves produce alcohol solutions that domestic goats (Dayton 1931)were reported to fluoresce brownish-red during exposure to intensively graze black sagebrush. Use may long-wave ultraviolet light (Winward 1980). depend on the variety, form, or subspecies present (Brunner 1972). Cultural treatment Vegetation and seeding of forage or cover plants are not Plant species associated with black sage- likely to improve composition and structural brush communities are often the same as those qualities of black sagebrush stands. Disturb- found with stiff and low sagebrush communi- ance of these stands initiates long succession ties. Species lists from black sagebrushlbunch- from Sandberg bluegrass dominance back to grass stands regularly include one or more of other bunchgrasses. Often, stands in poor condition have not recovered although they were the following: bearded bluebunch wheatgrass, protected from grazing (Winward 1980). Indian ricegrass, bottlebrush squirreltail, and Sandberg bluegrass; occasionally, needleandthread may be present (Winward 1980). 22-EARLY LOW SAGEBRUSH/ Short stature and wide spacing of shrubs BUNCHGRASS throughout the stands create sufficient cover Early low sagebrush has also been called for hiding or thermally protecting only the alkali sagebrush (Beetle 1960, Tisdale et al. smallest animals (table 8). 1969) and early sagebrush (Winward 1980). This species is also found primarily east of Site Oregon, but limited local stands exist in Vegetation stands of black sagebrush/ Deschutes, Crook, Lake, and Harney Counties bunchgrass occur between 1 200 and 2 740 (Beetle 1960, Brunner 1972, Tisdale et al. 1965, meters (4,000 and 9,000 ft) on drier, shallow Winward 1980). Eckert (1957)reported a large- sites (Beetle 1960, Dayton 1931, Winward headed form of low sagebrush that flowered

47 from mid-July until mid-August in northern fine textured and highly alkaline soils (Beetle Lake and Harney Counties; the phenology of 1960). In Idaho, Tisdale et al. (1965)associated the form he observed matches that of this this species with a silty clay loam subsoil at 0.1 uniquely early-flowering species of short meter (0.3 ft) and a strongly developed B sagebrush. horizon at 0.3-0.4 meter (1-2 ft).

The form and stature of early low sage- Discussion brush and the sites it is common to in many ways resemble those of low sagebrush. It is a Early low sagebrush, except for its earlier dwarf shrub (fig. 21C) with lax and spreading flowering, may be best considered a form of branches that frequently layer (Beetle 1960). low sagebrush (Winward 1980). Its preference These two short sagebrushes, early low and as forage by mule deer has not been evaluated. low, form adjacent pure stands; but they may This species was not grazed in Nevada and was also occur mixed together (Eckert 1957, Win- seldom eaten by sage grouse (Brunner 1972). ward 1980). These species cannot be readily Management of early low sagebrush stands separated on the basis of leaf form or should enhance or preserve the diversity of fluorescence. Alcohol extract solutions fluor- native forage species and adjust the grazing esce creamish-blue from exposure to long-wave season to accomplish that goal. Crested wheat- ultraviolet light (Winward 1980). Difference in grass is not adapted to the sites because of the phenology is the most suitable diagnostic shallow, heavy, and seasonally saturated soils. characteristic to separate early low and low sagebrush (see footnote 8, p 36). Other Commu nities Vegetation by Shrubs Plants associated with early low sagebrush are commonly found in stands of other short 23-SQUAW APPLElBUNCHGRASS (fig. 33) sagebrushes. Typical lists include pussytoes, Squaw apple is an intricately and rigidly wild buckwheat, phlox, violet, Idaho fescue, branched, deciduous shrub that may attain Thurber needlegrass, Sandberg bluegrass, and heights of 2 meters (6.5 ft) (Hitchcock and bottlebrush squirreltail (Tisdale et al. 1965). Cronquist 1973). Subspecies or varieties ha.ve Early low sagebrush stands offer little not been identified. structural cover because of plant density (table 8).Stands rich in species may serve as valuable forage areas, but only small animals find much shelter in typical stands.

Site Early low sagebrush stands are found at elevations from 1 830 to 2 440 meters (6,000 to 8,000 ft) on poorly drained, hot, and dry sites; the ecological situation is similar to sites dominated by low sagebrush (Beetle 1960, Winward 1980.)

Soil Brunner (1972) found early low sagebrush Figure 33.-Squaw applelbunchgrass corn- in pure stands on shallow stony soils, but the munity illustrating the large, robust form species also occurred in mixed stands with of the species and other species that corn- other sagebrushes on deeper soils. The name monly occur with it, such as mountain big “alkali sagebrush” came from plants found on sagebrush and bearded bluebunch wheatgrass (Chris Maser photograph).

48 Vegetation ally characterized by rolling hills and prominent drainages. Although squaw apple is seldom the dominant (measured by canopy cover) shrub of Soi I broad expanses, it may occasionally assume dominance in some stands. Squaw apple con- Squaw apple commonly occurs on stitutes the tallest shrub layer except where moderately deep, well-drained soils. Texture antelope bitterbrush is present. A variety of generally varies from loam to sandy loam. The other shrub species comprise the shorter shrub percent of soil surface covered by cobbles and layer. Squaw apple also occurs in eastern stones and cobble and stone volume in the soil Oregon in association with scattered pon- profile are highly variable. derosa pine and western juniper. Squaw apple also occurs on very shallow, Squaw apple communities may have as medium-textured soils derived from weakly many as four major strata, the tallest at a consolidated, occasionally fractured tuff and height of 1.5 meters (5 ft). Mountain big diatomaceous deposits. Plant density is likely sagebrush may average 0.8 meter (2.5 ft)and is controlled, in part, by fracturing frequency of usually the dominant shrub of the middle the soil parent material. Plant crowns act as a shrub layer with green and gray rabbitbrush protective influence on these highly erodible occasionally present. The subshrub granite soils." gilia and the shrublike slenderbush eriogonum comprise a sparse, low shrub layer 0.5 meter Discussion (1.5 ft) tall. The herbaceous layer averages 0.2 Hayes and Garrison (1960) described meter (0.5 ft) tall and consists primarily of wild squaw apple as fair browse for domestic sheep buckwheat, fleabane, rockcress, low pussytoes, and cattle in spring and for deer in the winter. Thurber needlegrass, bearded bluebunch Beneath the canopy of squaw apple is a well- wheatgrass, Sandberg bluegrass, and cheat- concealed haven for small animals. Stem den- grass. sity is high in the interior of squaw apple Communities are characterized by a shrub crowns, providing small birds excellent protec- cover of approximately 10-15 percent. tion from predators. Often two to five squaw apple bushes may be found growing close together so that their 24-BLACK GREASEWOOD/GRASS (fig. 34) branches intertwine, giving the illusion of one Black greasewood is a spiny, freely huge shrub, 2.0-2.5 meters (6.5-8 ft) in branched shrub that may grow" nearly 2.5 diameter. meters (8 ft) high (Hitchcock and Cronquist Threetip sagebrush, averaging 0.6 meter (2 1973). The succulent leaves of this deciduous f t) tall, occasionally dominates the inter- species give a luxuriant appearance to the mediate shrub layer. Also included in this layer plant (Shantz and Piemeisel 1940). The bright yellow green of communities dominated by are a few scattered big sagebrush, gray rab- black greasewood contrasts with the gray bitbrush, and green rabbitbrush. Threetip green of sagebrush communities during the sagebrush and the rabbitbrushes are rounded summer. The species is not relished by shrubs that branch from near the base. The livestock, but severe grazing may produce basal leaves of wild buckwheat and low pussy- smaller plants with compact canopies. Black toes may form small islandlike mats on the greasewood will resprout from the root crown otherwise stony and bare surface. after fire (Daubenmire 1970).

Site "J. Michael Geist, unpublished data on file at Range and Wildlife Habitat Laboratory, La Grande, Oregon The range of squaw apple is restricted to high, moist hillsides in the northern portion of southeastern Oregon. The topography is gener-

49 both in percent composition and height-1.3 meters (4.3 ft). Big sagebrush and green rabbitbrush also occur. In some communities, giant wildrye attains the greatest height of any species but is present only in small amounts; however, this grass may have been more preva- lent before grazing. A layer of cheatgrass is common between and beneath the shrubs, and the amount of bare ground varies according to fluctuations in the density of this species.

Site The black greasewoodlgrass community reaches its best development on lowlands in Figure 34.-The black greasewoodlgrass the Great Basin of southeastern Oregon, occur- community is one of the tallest of the ring primarily on saline-sodic flood plains, shrub types and provides excellent protec. tive cover for nesting birds and small mam- playas, and terraces. On middle slopes of about mals and thermal cover for all species. 10 percent, spiny hopsage, as well as giant wildrye and green rabbitbrush, is conspicuous Vegetation in the black greasewood community. On the lower slopes, spiny hopsage and shadscale Generally, black greasewood communities saltbush become prominent. The low growing have shrub and herbaceous layers, but com- bud sagebrush occurs sparingly, and cheat- munity structure varies, depending on species grass dominates the herbaceous layer. composition or age class. Some black greasewood areas contain few Soi I other shrubs, and in pure stands the plants are widely spaced with large interspaces of bare The black greasewoodlgrass community soil (Shantz and Piemeisel 1940).On some sites generally occurs on soils exhibiting high with almost pure stands, the shrub cover alkalinity and high water tables (Potter 1957); ranges from 10 to 20 percent and averages be- however, Shantz and Piemeisel (1940)reported tween 0.9 meter and 1.2 meters (3and 4 ft) tall. alkali is not necessary for the presence of black The interspaces are mainly barren but are occa- greasewood, but a high moisture content of the sionally occupied by cheatgrass and bottle- soil is. brush squirreltail. Russianthistle is present Discussion but restricted to disturbed areas near roads. The black greasewoodlgrass community Black greasewood comprises 90-95 percent provides thermal cover for all species of of the shrubs in stands of the valley bottoms; wildlife and excellent protective cover for the remainder consists of very large shadscale nesting birds and small mammals. This excel- saltbush plants (reaching heights of 1 meter- lent cover is primarily due to the tall growth 3.28 ft), green rabbitbrush, and basin big form of black greasewood combined with its sagebrush. Cheatgrass, which usually spines and coarse structure: dominates the herbaceous layer, is confined beneath shrub canopies. Rickard (1964) reported sagebrush was replaced by black greasewood in an area that Patches of the strongly rhizomatous desert had been excessively grazed. The attendant saltgrass occur in areas of greater moisture. loss of a relatively nonsaline surface layer pro- Desert saltgrass swards occur in some dense duced a saline-sodic environment where water black greasewood communities where the was available only to the halophytic black shrub cover may reach almost 30 percent. greasewood. This indicates the limited options Black greasewood dominates the shrub layer,

50 available for management of stands that occur Average height of the grass-forb layer in on marginal sites. this community is 14 centimeters (6 inches), ranging from 5 to 25 centimeters (2 to 10 25-SHADSCALE SALTBUSH/ inches). In most cases, total crown cover for BUNCHGRASS (fig. 35) the forb component is less than 1 percent (see footnote 5, p. 18). Dominance of the her- Vegetation baceous layer varies between Sandberg blue- Shadscale saltbusWbunchgrass is the grass and bottlebrush squirreltail; average dominant community on large expanses of the crown cover of each species is generally less high desert in southeastern Oregon (see foot- than 5 percent. Sandberg bluegrass occurs in note 5, p. 18). Crown cover in most stands shrub interspaces and under shrub crowns; averages 15 percent and ranges from 10 to 35 bottlebrush squirreltail, however, is found pri- percent. Heights average 37 centimeters (15 marily under shrubs. The average height of inches) and range from 30 to 45 centimeters (12 Sandberg bluegrass, including seedstalks, is 10 to 18 inches). Spiny hopsage is a common centimeters (4 inches); that of bottlebrush species, generally with a crown cover of less squirreltail, 15 centimeters (6 inches). Cheat- than 5 percent; height averages 62 centimeters grass, the only other grass present, has an (24 inches) and ranges from 50 to 70 centi- average crown cover of less than 1 percent, meters (20 to 28 inches) between sites. Big Forbs are sparse in this type. Only clasping sagebrush is. a sparse component in most pepperweed and Russianthistle are common. areas, with an average crown cover of less than Crown cover is generally less than 5 percent 5 percent and an average height of 56 centi- and rarely exceeds 15 percent. Rare species are meters (22 inches), ranging from 50 to 70 centi- halogeton, wild buckwheat, and milkvetch. meters (20 to 28 inches). Big sagebrush generally appears as small patches within the type, Site indicating a possible variation in ‘soil. Bud sagebrush is common on most sites, with a Most stands of this community occur on crown cover averaging less than 5 percent but relatively level, lowland sites, but some are ranging up to 10 percent in some areas; present on rolling topography. Slope varies average height is 15 centimeters (6 inches), from 0 to 30 percent among sites, directional ranging from 10 to 20 centimeters (4 to 8 aspect from north-northwest to south; the com- inches). The only other shrubs commonly pres- munity is found on all slope positions. Macro- ent are gray and green rabbitbrushes, normally relief is level to moderately rolling hills, and with less than 1-percent crown cover. Heights microrelief is moderately smooth to moder- of gray and green rabbitbrushes average 48 ately rough (see footnote 5, p. 18). and 30 Centimeters (19 and 12 inches), - respectively. Soil Soils are moderately deep, varying from silt loams to coarse gravelly and stony silt loams. Cobble and stone cover of the soil surface varies from 5 to 40 percent, with erosion pavement made up of coarse gravels that occupy small, low areas in shrub interspaces. In some lowland flats, dry soil surfaces in depression spots exhibit cracked and plated, polygonal patterns. Soil in all areas is strongly alkaline. In most areas, edges and bottoms of surface stone are encrusted with a thin calcareous layer (see footnote 5, p. 18). Figure 35.-Shadscale salt bushlbunchgrass community occurs on large expanses of the high deserts (Chris Maser photograph).

51 Discussion Grassland Communities Vegetation in all stands appears to have had a long period of heavy use by livestock. The following natural communities Most perennial bunchgrass plants are present dominated by grass and grasslike plants in- only under the protection of shrub crowns. The clude seasonally wet and wet meadows, as well most common forbs are annual species (Rus- as subalpine grasslands. Shrubs and trees are sianthistle and clasping pepperweed) on either minor components or are absent from abused ranges. Rodent burrows, mostly of these communities. kangaroo rats, are present in most sites but concentrated in areas of rolling, hilly relief. 26-SEASONALLY WET MEADOWS Mounds in these areas are large, 2 to 5 meters (figs. 36 and 37) (7 to 16 ft) in diameter, and occur on ridges and Vegetation slopes at a density up to 25 per hectare (10 per acre). Fewer and smaller mounds are found in At low elevations, seasonally wet meadow bottoms and on flat topography. Small vegetation is comprised primarily of grasses mounds of burrowing rodents are present on and sedges. Shrubs, such as willow, may occur level areas with a density up to 12 per hectare in riparian situations adjacent to the type, or (5 per acre). Large mounds vary from 4.5 to 6 dry site shrubs may be present in stands on meters (15 to 20 ft) in diameter and occur on high ground or upslope at the dry edge of the flats at a density of approximately 10 per hec- type. Grasses include Kentucky bluegrass, tare (4 per acre). Sandberg bluegrass, skyline bluegrass, Wheeler bluegrass, northern meadow barley, The primary dispersal pattern of halogeton and spike trisetum. Sedges that may be pres- into the shadscale saltbush type in the Great ent are smallwing, epapiilous, Douglas, Basin of Oregon is from highway shoulders Liddon, slenderbeak, and Raynolds. Rushes and barrow pits to rodent mounds and other generally occur as minor species; the most disturbed areas. From a survey of two 300- common in southeastern Oregon may be meter (985-ft) by 10-meter (33-ft) transects, Drummond and swordleaf (Hansen 1956). beginning 10 meters from a highway shoulder and moving at a right angle to the road, only 2 Hansen (1956) described shrubs in sub- halogeton plants were discovered in vegetation alpine seasonally wet meadows on Steens undisturbed by rodents, and 12 plants or small Mountain as low weatherbeaten forms typical groups of plants were counted on rodent of windswept high mountain vegetation. Com- mounds. Frequency of halogeton fell off with mon shrubs (willows listed in appendix under distance from the highway. “TREES”) are Drummond willow, coyote willow, snow willow, bush cinquefoil, cartha- Holmgren and Hutchings (1972) described moid goldenweed, and dwarf blueberry. Only shadscale saltbush as being palatable to four grass and grasslike species were re- livestock, but because of its spiny stems, use ported-false-scirpus sedge, subnigricans was limited to 15 to 20 percent of the previous sedge, winter bentgrass, and spreading wood- season’s growth. Bud sagebrush was generally rush. Forbs were more abundant: crag aster, palatable only in late winter, but it may be con- elkslip marshmarigold, alpine shootingstar, sumed all winter on occasion. It is considered Sierra willow weed, Watson willow weed, whit e more palatable for domestic sheep than for bogorchid, oblongleaf bluebells, Rocky Moun- cattle. Russianthistle and cheatgrass are often tain parnassia, lousewort, American bistort, abundant on heavily grazed shadscale saltbush Kellogg ’s knotweed, shor t-leaved cinquefoil, ranges. These species are considered both plainleaf buttercup, modest buttercup, rose- palatable and nutritious by Holmgren and root stonecrop, and crisped starwort. Hutchings (1972) but are undependable sources of forage because they are annuals.

52 Figure 36.-Extensive seasonally wet meadows are shown with lesser amounts of permanently wet meadows along streams and around ponds.

Figure 37.-Seasonally ponded sumps at the western edge of the Great Basin generally dry up by mid- season. These areas are severely limited in production of forage.

53 Site 27-PERMANENYLY WET MEADOWS (fig. 38) Seasonally wet meadows can occur in any vegetation zone in the high desert, from Vegeta tion bottomlands of low elevation to subalpine Plant species composition of stands varies ranges. Communities are present as either considerably among permanently wet sites on seasonally ponded or wet swales with no exter- rangelands of the high desert. Grasses and nal drainage or as transitional stands between sedges are typically dominant in meadows in wet meadows and dryland shrub types (Dealy good condition, and the forbs may also be 1971). Where streams have been channeled because of erosion, wet meadows often have abundant. At elevations below the subalpine been permanently converted to either season- zone, Hansen (1956) stated that whiplash ally wet meadows or dryland shrub communi- willow was the most abundant shrub in ties. In high precipitation zones or on sites meadows. Common grass and grasslike species where a subsurface lateral drainage emerges at were creeping bentgrass, northern meadow valley edges (toe slopes), the seasonally wet barley, mountain hairgrass, and ovalhead grassland community often becomes promi- sedge. Columbia monkshood, primrose mon- nent. Seasonal seeps, springs, and streams in keyflower, and Scouler St. Johnswort were the any slope position often provide small stands most common forbs. of the community.

Soi I Seasonally wet meadows generally occur on moderately deep to deep clay or clay loam soils. Soils under meadows in good condition are high in organic matter and moisture- holding capacity. Wide cracks occur in the dry clay soils, producing a churning action (surface soil falls into cracks when the soil is dry and is incorporated at a lower level when soils become wet and cracks close) which mixes the upper horizons. Trampling of meadows by livestock during wet periods can result in soil damage Figure 38.-Small areas of permanently wet and thus damage forage plants. meadows around seeps and springs are typical of the high desert mountains. Discussion Site Seasonally wet meadows are highly attrac- tive to domestic and wild ungulates, and heavy Other than the large meadowlands on grazing for more than 100 years has depleted private holdings, wet meadow communities are vegetation of these stands. Sites large enough found primarily as belts adjacent to riparian to provide substantial forage may be practical zones and as patches around perennial springs for intensive management of livestock. Small and seeps. They can be found on any aspect sites, however, such as near seasonal seeps and and most slopes, but those around springs and springs, may be best developed for wildlife but seeps may be more common on northerly with water piped outside the protected habitat. aspects. Young tender forbs and grasses available into late spring contribute to optimum habitat for Discussion young grouse (Bump et al. 1947) and probably Heavy grazing of most wet meadows for other gallinaceous birds. An oasis such as this over 100 years has had a marked effect on the is used by many wildlife species for food and composition and productivity of these com- cover. munities. Through recently developed manage-

54 ment practices, some riparian zones and adjacent wet meadows are being improved.12Small areas of meadow around springs and seeps within expanses of drier range may appear to be uneconomical for management planning, yet they are stands where livestock and wildlife concentrate. Because of their special values for wildlife, it may be desirable to protect them to enhance the production of grasses, forbs, and shrubs and thus improve cover and forage for a wide variety of small animals and birds.

28-SUBALPIN E BUNCHGRASS (figs. 39 and 40) Figure 39.-Subalpine grassland of the high desert mountains; sheep fescue domi- Vegetation nates the site (Chris Maser photograph). Grassland communities of the subalpine zone are characterized by a dominant stand of fescue, primarily sheep fescue (Maser and Strickler 1978). Other grasses and grasslike species are rough fescue, prairie junegrass, Cusick bluegrass, longtongue mutton bluegrass, Sandberg bluegrass, and Dunhead sedge (Hansen 1956). Areas with disturbed soil contain small amounts of forbs (Hansen 1956).

Site Hansen (1956) found the subalpine zone above 2 439-meter (8,000-ft)elevation predominantly grassland interspersed with large areas of bare windswept ground and the lower 244 meters (800 ft) overgrown to short sagebrush. Figure 40.-Detailed view of a subalpine grassland; sheep fescue dominates the Discussion site (Chris Maser photograph). Communities in subalpine regions are sheep during the peak of the grazing season. fragile and can be severely damaged by live- Mule deer, pronghorns, bighorn sheep, and stock grazing and trampling. High winds and occasionally Rocky Mountain elk also use unstable soils combined with severe grazing these high elevation areas. can produce rapid erosion of the surface layer of grassland soils. Large, bare windswept areas are probably a product of severe use by Successional Vegetation domestic sheep during the late 1800’s and early 1900’s. Griffiths (1902) toured through Successional patterns are highly complex southeastern Oregon and referred to domestic in the communities of the northern Great sheep use as extremely heavy. Vegetation was Basin. Current composition of plant species severely damaged. Griffiths (1902)noted large can be misleading when site potential and clouds of dust caused by bands of domestic suitable management alternatives are determined for various stands. Because of altera- 12Persona1communication, 1979, from Robert Kindschy, Bureau of Land Management, Vale, Oregon. tions by natural fire patterns and a long

55 history of heavy use by livestock, there are some stage of succession. Nearly complete relatively few communities or local stands in destruction of the grass-forb layer under a the desert rangelands that can be considered at shrub or tree-shrub-dominated stand can also or near climax condition. In other words, few make identification of plant communities dif- stands are in a natural balance with the envi- ficult. Here, discovery of remnants of grass ronment. The stands most difficult to identify species and an analysis of site factors can help are those that have been most severely altered. identify the original plant community. Much work has been done to identify An important key to developing a manage- subspecies of big sagebrush (Beetle 1960, ment outlook is recognizing ecological poten- Beetle and Young 1965, Winward 1970, tial of the site-the potential of a community Winward and Tisdale 1977). Some of this work to support vegetation of interest. For example, includes identification of typical soils and sites Burkhardt and Tisdale (1969)presented strong on which various subspecies are found. For evidence that with management and fire con- example, Winward and Tisdale (1977), besides trol in southwestern Idaho, western juniper giving a taxonomic description of the sub- has the potential to occupy the entire moun- species, describe basin big sagebrush as tain big sagebrush/bunchgrass community. growing “in deep, well drained soils of valley Given a site with this potential, a seed source, bottoms and lower foothill regions.” They and barring fire or other repression of the trees describe the occurrence of the subspecies in and shrubs, the inevitable conclusion will be a terms of elevation and response to disturbance. western juniperlbig sagebrushlbunchgrass Recognition of subspecies and information on stand. Similarly, given a potential sagebrush/ sites, along with soils and identification of bunchgrass community in a bunchgrass suc- remnant species that are characteristic of cessional stage, the inevitable conclusion, climax or advanced successional communities, barring inhibition of developing sagebrush, can often help managers determine potential will be a shrublgrass community. With such for a community. knowledge built into management plans, managers will be spared the frustration of see- The relationship between dominance of ing “invasions” of shrubs or trees into com- sagebrush and that of bunchgrass in climax munities considered stable “grass” types or shrub steppe communities is not understood. “shrublgrass” types. The managers can then An excellent stand of grass dominated by plan management strategies of stocking rates, bearded bluebunch wheatgrass or Idaho fescue season of use, manipulation of vegetation, and and bearded bluebunch wheatgrass, either “preserved diversity” to suit established devoid of big sagebrush or having a small priorities based on the ecological realities of component of this shrub, may appear on the each site’s potential. surface as a climax grassland community. That premise is usually not the case in the The rate of successional advancement can shrub steppe. Central and southeastern vary greatly within any community. Expe- Oregon steppe communities devoid of a major rienced managers generally recognize and have shrub component are uncommon, and those a feel for this. A light fire that leaves scattered that do appear as grasslands may have re- shrubs in a sagebrush type provides a ready sulted from burning (Franklin and Dyrness seed source for reoccupancy by shrubs. A hot 1973). extensive fire, on the other hand, may virtually eliminate seed sources of shrubs over a great A brief examination of sites in question distance. Burning a south exposure shrub/ should help observers to correctly classify the stands. If topography, elevation, and soils in grass community can produce microclimate areas of question are similar to adjacent types conditions that may prevent reestablishment of shrubs for a long time. Such effects can be with stands of shrubland communities, there is assessed and the information used in project- a greater than 90-percent chance that the ing future management alternatives. questionable type is a shrub-grass climax in

56 Literature Cited Cook, C. W. 1972. Comparative nutritive values of Adams, A. W. forbs, grasses, and shrubs. In Wildland 1975. A brief history of juniper and shrub shrubs: their biology and utilization. C. M. populations in southern Oregon. Oreg. McKell, J. P. Blaisdell, and J. R. Goodin, Wildl. Comm., Wildl. Res. Rep. 6, 33 p. eds. USDA For. Serv. Gen. Tech. Rep. Corvallis . INT-1, p. 303-310. Intermt. For. and Range Alderfer, J. M. Exp. Stn., Ogden, Utah. 1976. A taxonomic study of bitterbrush Cornelius, D. R., andM. W. Talbot. (Purshia tridentata (Pursh) DC.) in Oregon. 1955. Rangeland improvement through M.S. thesis. Oreg. State Univ., Corvallis. seeding and weed control on east slope 197 p. Sierra Nevada and on southern Cascade Anderson, E. W. Mountains. U.S. Dep. Agric. Agric. Handb. 1956. Some soil-plant relationships in 88,51 p. eastern Oregon. J. Range Manage. Culver, R. N. 9(4):17 1-1 75. 1964. An ecological reconnaissance of the Anthony, H. E. Artemisia steppe on the east central 1928. Field book of North American mam- Owyhee uplands of Oregon. M.S. thesis. mals. 674 p. G. P. Putnam’s Sons, New Oreg. State Univ., Corvallis. 99 p. York. Daubenmire, R. Beetle, A. A. 1956. Climate as a determinant of vegeta- 1960. A study of sagebrush, the section tion distribution in eastern Washington and Tridentatae of Artemisia. Agric. Exp. Stn. northern Idaho. Ecol. Monogr. Bull. 368,83 p. Univ. Wyo., Laramie. 26(2):131 -154. Beetle, A. A., and A. Young. Daubenmire, R. 1965. A third subspecies in the Artemisia 1968. Plant communities, a textbook of triden tata complex. Rhodora 67:405-406. plant synecology. 300 p. Harper and Row, Bella, D. A., and W. S. Overton. Publ., New York. 1972. Environmental planning and eco- Daubenmire, R. logical possibilities. J. Sanitation Eng. Div., 1970. Steppe vegetation of Washington. Proc. Am. SOC.Civ. Eng. 98(SA3). Wash. Agric. Exp. Stn. Tech. Bull. 62, Brunner, J. R. 135 p. Wash. State Univ., Pullman. 1972. Observatims on Artemisia in Dayton, W. A. Nevada. J. Range Manage. 25(3):205-208. 1931. Important western browse plants. Bump, G., R. W. Darrow, F. C. Edminster, and U.S. Dep. Agric. Misc. Publ. 101, 214 p. W. F. Crissey. Washington, D.C. 1947. The ruffed grouse - life history, Dealy, J. E. propagation, and management. 915 p. N. Y. 1971. Habitat characteristics of the Silver State Conserv. Dep., Buffalo. Lake mule deer range. USDA For. Serv. Burkhardt, J. W., andE. W. Tisdale. Res. Pap. PNW-125, 99 p. Pac. Northwest 1969. Nature and successional status of For. and Range Exp. Stn., Portland, Oreg. western juniper vegetation in Idaho. J. Dealy, J. E. Range Manage. 22(4):264-270. 1975. Ecology of curlleaf mountain- Burkhardt, J. W., andE. W. Tisdale. . mahogany in Oregon and adjacent areas. 1976. Causes of juniper invasion in Ph. D. thesis. Oreg. State Univ., Corvallis. southwestern Idaho. Ecology 57(3):472-484. 162 p. Concannon, D. M. a Dealy, J. E., and J. M. Geist. 1978. Plant succession on burned areas of 1978. Conflicting vegetational indicators the A r t e m isia t riden ta ta/Agrop y ro n on some central Oregon scablands. J. Range spicatum habitat type in southeastern Manage. 31(1):56-59. Oregon. M.S. thesis. Humboldt State Univ., Arcslta, Calif. 101 p.

57 Dealy, J. E., J. M. Geist, and R. S. Driscoll. Fosberg, M. A., and M. Hironaka. 1978a. Communities of western juniper in 1964. Soil properties affecting the distribu- the intermountain Northwest. In Pro- tion of big and low sagebrush communities ceedings of the western juniper ecology and in southern Idaho. Am. SOC.Agron. Spec. management workshop. R. E. Martin, J. E. Publ. 5~230-236. Dealy, and D. L. Caraher, eds. USDA For. Franklin, J. F., and C. T. Dyrness. Serv. Gen. Tech. Rep. PNW-74, p. 11-29. 1973. Natural vegetation of Oregon and Pac. Northwest For. and Range Exp. Stn., Washington. USDA For. Serv. Gen. Tech. Portland, Oreg. Rep. PNW-8, 417 p. Pac. Northwest For. Dealy, J. E., J. M. Geist, and R. S. Driscoll. and Range Exp. Stn., Portland, Oreg. 197813. Western juniper communities on Gabrielson, I. N., and S. G. Jewett. rangelands of the Pacific Northwest. Proc. 1940. Birds of Oregon. 650 p. Oreg. State 1st Int. Rangeland Congr., p. 201-204. D. N. Coll. [Univ.] Publ., Corvallis. Hyder, ed. SOC.Range Manage., Denver, Garrison, G. A., A. J. Bjugstad, D. A. Duncan, Colo. M. E. Lewis, and D. R. Smith. Dean, M. L. 1977. Vegetation and environmental fea- 1960. A taxonomic and ecologic study of tures of forest and range ecosystems. U.S. the vascular plants of a section of the Dep. Agric. Agric. Handb. 475, 68 p. Owyhee River Canyon in Oregon. M.S. Washington, D. C. thesis. Oreg. State Univ., Corvallis. 147 p. Garrison, G. A., J. M. Skovlin, C. E. Poulton, deBoer, T. A. and A. H. Winward. 1974. [The contribution of vegetation 1976. Northwest plant names and symbols science to the management of wildlife for ecosystem inventory and analysis. reserves.] Misc. Pap. 5, p. 63-78. Land- USDA For. Serv. Gen. Tech. Rep. PNW-46, bouw hogesc h. W ageningen, The Nether- 263 p. Pac. Northwest For. and Range Exp. lands. [In Dutch.] Stn., Portland, Oreg. Dietz, D. R., R. H. Udall, andL. E. Yeager. Griffiths, D. 1962. Chemical composition and digestibil- 1902. Forage conditions on the northern ity by mule deer of selected forage species, border of the Great Basin. USDA Bur. Plant Cache La Poudre Range, Colorado. Colo. Ind. Bull. 15, 60 p. Gov. Print. Off., Game and Fish Dep. Tech. Publ. 14,89p. Washington, D.C. Hall, F. C. Dietz, D. R., and L. E. Yeager. 1973. Plant communities of the Blue Moun- 1959. The apparent role of sagebrush in the tains in eastern Oregon and eastern management of mule deer winter range. Washington. USDA For. Serv. R6 Area Proc. West. Assoc. State Game and Fish Guide 3-1, 62 p. Pac. Northwest Reg., Comm. 39:151-158. Portland, Oreg. Driscoll, R. S. Hanks, D. L., J. R. Brunner, D. R. Christensen, 1964. Vegetation-soil units in the central and A. P. Plummer. Oregon juniper zone. USDA For. Serv. Res. 1971. Paper chromatography for determin- Pap. PNW-19, 60 p. Pac. Northwest For. ing palatability differences in various and Range Exp. Stn., Portland, Oreg. strains of big sagebrush. USDA For. Serv. Eckert, R. E., Jr. Res. Pap. INT-101, 9 p. Intermt. For. and 1957. Vegetation-soil relationships in some Range Exp. Stn., Ogden, Utah. Artemisia types in northern Harney and Lake Counties, Oregon. Ph. D. thesis. Oreg. Hanks, D. L., E. D. McArthur, R. Stevens, and State Univ., Corvallis. 208 p. A. P. Plummer. Eckert, R. E. 1973. Chromatographic characteristics and 1958. Vegetation-soil relationships in some phylogenetic relationships of Artemisia Artemisia types in Harney and Lake Coun- tridentata. USDA For. Serv. Res. Pap. INT-141, 24 p. Intermt. For. and Range ties, Oregon. J. Range Manage. 11(1):50. Exp. Stn., Ogden, Utah.

58 Hansen, C. G. Kuchler, A. W. 1956. An ecological survey of the 1964. The potential natural vegetation of vertebrate animals on Steen’s Mountain, the conterminous United States. Am. Harney County, Oregon. Ph. D. thesis. Geogr. SOC.Spec. Pubi. 36,160 p. Oreg. State Univ., Corvallis. 200 p. Kufeld, R. C., 0. C. Wallmo, and C. Feddema. Hayes, D. W., andG. A. Garrison. 1973. Foods of the Rocky Mountain mule 1960. Key to important woody plants of deer. USDA For. Serv. Res. Pap. RM-111, eastern Oregon and Washington. U.S. Dep. 31 p. Rocky Mt. For. and Range Exp. Stn., Agric. Agric. Handb. 148, 227 p. U.S. Gov. Fort Collins, Colo. Print. Off., Washington, D.C. Leckenby, D. A. Hill, R. R. 1977. Management of mule deer and their 1956. Forage, food habits, and range habitat: Applying concepts of behavior, management of the mule deer. In The deer of physiology, and microclimate. Proc. West. North America, p. 393-414. W. P. Taylor, ed. Assoc. State Game and Fish Comm. Stackpole Co., Harrisburg, Pa. 57 ~206-217. Hitchcock, C. L., and A. Cronquist. Leckenby, D. A. 1973. Flora of the Pacific Northwest, an 1978a. Mule deer occupancy of plant com- illustrated manual. 730 p. Univ. Wash. munities on a south-central Oregon winter Press, Seattle. range. Spec. Rep., Oreg. Dep. Fish and Hitchcock, C. L., A. Cronquist, M. Ownbey, Wildl. Final Rep. Proj. W-53-R and W-70-R- and J. W. Thompson. Sub. B. SIJ5.77 p. 1955. Vascular plants of the Pacific North- Leckenby, D. A. west. Part 5: Compositae. 343 p. Univ. 1978b. Western juniper management for Wash. Press, Seattle. mule deer. In Proceedings of the western Hitchcock, C. L., A. Cronquist, M. Ownbey, juniper ecology and management workshop. and J. W. Thompson. R. E. Martin, J. E. Dealy, andD. L. Caraher, 197.7. Vascular plants of the Pacific North- .eds. USDA For. Serv. Gen. Tech. Rep. west (5 parts). Part 3, Saxifragaceae to PNW-74, p. 137-161. Pac. Northwest For. Ericaceae. 614 p. Univ. Wash. Press, Seattle and Range Exp. Stn., Portland, Oreg. and London. Leckenby, D. A., and D. E. Toweill. Holmgren, R. C., andS. S. Hutchings. 1979. Mule deer winter range ecology and 1972. Salt desert shrub response to grazing management: Seeding response of selected use. In Wildland shrubs: Their biology and plant species on a mule deer winter range in utilization. C. M. McKell, J. P. Blaisdell, south-central Oregon. Oreg. Dep. Fish and and J. R. Goodin, eds. USDA For. Serv. Wildl. Final Rep. W-53-R-12 and W-70-R-1 Gen. Tech. Rep. INT-1, p. 153-164. Intermt. through 8, Sub. B. SIUJ2.22 p. For. and Range Exp. Stn., Ogden, Utah. MacArthur, R. H., and J. W. MacArthur. Hormay, A. L. 1961. On bird species diversity. Ecology 1943. Bitterbrush in California. USDA For. 42(3):594-598. Serv., Calif. For. and Range Exp. Stn. Res. McIlvain, E. H., andM. C. Shoop. Note 34,13 p. 1971. Shade for improving cattle gains and Ingles, L. G. rangeland use. J. Range Manage. 1967. Mammals of the Pacific States. 506 p. 24(3):181-184. Stanford Univ. Press., Stanford, Calif. Maser, C., and J. S. Gashwiler. Kornet, C. A. 1978. Interrelationships of wildlife and 1978. Status and habitat use of California western juniper. In Proceedings of the bighorn sheep on Hart Mountain, Oregon. western juniper ecology and management M.S. thesis. Oreg. State Univ., Corvallis. workshop. R. E. Martin, J. E. Dealy, and D. L. Caraher, eds. USDA For. Serv. Gen. 49 p. ’ Kuchler, A. W. Tech. Rep. PNW-74, p. 37-82. Pac. North- 1957. Applied phytosociology. Ecology west For. and Range Exp. Stn., Portland, 38(3):541-542. Oreg.

59 Maser, C., and G. S. Strickler. Rickard, W. H. 1978. The sage vole, Lagurus curtatus, as 1964. Demise of sagebrush through soil an inhabitant of subalpine sheep fescue, changes. Bioscience 14(1):43-44. Festuca ouina, communities on Steens Rittenhouse, L; R., and F. A. Sneva. Mountain: An observation and interpreta- 1977. A technique for estimating big tion. Northwest Sci. 52(3):276-284. sagebrush production. J. Range Manage. Miller, A. H. 30(1):68-70. 1968. Amphispiza belli Nevadensis Roberts, R. W. (Ridgeway) northern sage sparrow. In Life 1975. Evaluating the impact of range treat- histories of North American cardinals, gros- ment on vegetation. Oreg. Wildl. Comm., beaks, buntings, towhees, finches, sparrows, Wildl. Res. Rep. 7,30 p. Corvallis. and allies, part 2, p. 1004-1013. A. C. Bent Schumaker, G. A., and C. L. Hanson. and 0. L. Austin, Jr., eds. Dover Publ., Inc., 1977. Herbage response after mechanical New York. and herbicide treatment of big sagebrush in Mueller-Dombois, D., and H. Ellenberg. southwest Idaho. U.S. Dep. Agric. Agric. 1974. Aims and methods of vegetation Res. Serv. W-46, 15 p. ecology. 547 p. John Wiley and Sons, New Segura-Bustamante, M. York. 1970. Ecology of bitterbrush (Purshia Nagy, J. G., and R. P. Tengerdy. tridentata (Pursh) DC.) in the Silver Lake 1968. Antibacterial action of essential oils deer winter range, Oregon. Ph. D. thesis. of Artemisia as an ecological factor. Appl. Oreg. State Univ., Corvallis. 342 p. Microbiol. 16(3):441-444. Shantz, H. L., and R. L. Piemeisel. Nord, E. C. 1940. Types of vegetation in the Escalante 1965. Autecology of bitterbrush in Cali- Valley, Utah, as indicators of soil condi- fornia. Ecol. Monogr. 35(3):307-334. tions. U.S. Dep. Agric. Tech. Bull. 713, Odum, E. P. 46 p. U.S. Gov. Print. Off., Washington, 1971. Fundamentals of ecology. 3d ed. D.C. 574 p. W. B. Saunders Co., Philadelphia, Pa. Sheehy, D. P. Odum, E. P. 1975. Relative palatability of seven 1979. Ecological importance of the riparian Artemisia taxa to mule deer and sheep. M.S. zone. In Strategies for protection and thesis. Oreg. State Univ., Corvallis. 147 p. management of floodplain wetlands and Short, H. L., R. M. Blair, andL. Burkart. other riparian ecosystems. R. R. Johnson 1972. Factors affecting nutritive values. In and J. F. McCormick, tech. coords. USDA Wildland shrubs: Their biology and utiliza- For. Serv. Gen. Tech. Rep. WO-12, p. 2-4. tion. C. M. McKell, J. P. Blaisdell, and J. R. Washington, D.C. Goodin, eds. USDA For. Serv. Gen. Tech. Packard, P. L. Rep. INT-1, p. 311-318. Intermt. For. and 1972. Pinus ponderosa in Malheur County, Range Exp. Stn., Ogden, Utah. Oregon. Madrono 21(5):298. Smith, A. D. Pechanec, J. F., A. P. Plummer, J. H. Robert- 1950. Sagebrush as a winter feed for deer. son, and A. C. Hull, Jr. J. Wildl. Manage. 14(3):285-289. 1965. Sagebrush control on rangelands. Smith, A. D. U.S. Dep. Agric. Agric. Handb. 277, 40 p. 1952. Digestibility of some native forages U.S. Gov. Print. Off., Washington, D.C. for mule deer. J. Wildl. Manage. Peck, M. E. 16(3):309-312. 1961. A manual of the higher plants of Soil Survey Staff. Oregon. 2d ed. 936 p. Binfords and Mort, 1951. Soil survey manual. U.S. Dep. Agric. Publ., Portland, Oreg. Agric. Handb. 18, 503 p. U.S. Gov. Print. Potter, L. D. 0 f f ., Washi ngt on, D .C . 1957. Phytosociological study of San Stevens, R., and E. D. McArthur. Augustin Plains, New Mexico. Ecol. 1974. A simple field technique for iden- Monogr. 27(1):113-136. tification of some sagebrush taxa. J. Range Manage. 27 (4):325-326.

60 Thomas, J. W., R. M. DeGraaf, and J. C. Urness, P. J. Mawson. 1966. Influence of range improvement prac- 1977. Determination of habitat require- tices on composition, production, and ments for birds in suburban areas. USDA utilization of Artemisia deer winter range in For. Serv. Res, Pap. NE-357, 15 p. North- central Oregon. Ph. D. thesis. Oreg. State east. For. Exp. Stn., Upper Darby, Pa. Univ., Corvallis. 183 p. Thomas, J. W., C. Maser, and J. E. Rodiek. Volland, L. A. 1978. Edges-their interspersion, resulting 1976. Plant communities of the central diversity and its measurement. In Pro- Oregon pumice zone. USDA For. Serv. R6 ceedings of the workshop on nongame bird Area Guide 4-2, 110 p. Pac. Northwest Reg., habitat management in the coniferous Portland, Oreg. forests of the Western United States. West, N. E., R. J. Tausch, K. H. Rea, and P. T. USDA For. Serv. Gen. Tech. Rep. PNW-64, Tueller. p. 91-100. Pac. Northwest For. and Range 1978. Taxonomic determination, distribu- Exp. Stn., Portland, Oreg. tion, and ecological indicator values of Thomas, J. W., H. C. Black, Jr., R. J. sagebrush within the pinyon- juniper Scherzinger, and R. J. Pedersen. woodlands of the Great Basin. J. Range 1979a. Deer and elk. In Wildlife habitats in Manage. 3 1(2):87-92. managed forests: The Blue Mountains of Westoby, M., and F. H. Wagner. Oregon and Washington. J. W. Thomas, ed. 1973. Use of a crested wheatgrass seeding U.S. Dep. Agric. Agric. Handb. 553, by black-tailed jackrabbits. J. Range p. 104-127. U.S. Gov. Print. Off., Manage. 26 (5):349 -35 2. Washington, D.C. Winward, A. H. Thomas, J. W., C. Maser, and J. E. Rodiek. 1970. Taxonomic and ecological relation- 197913. Riparian zones. In Wildlife habitats ships of the big sagebrush complex in Idaho. in managed rangelands: The Great Basin of Ph. D. thesis. Univ. Idaho, Moscow. 80p. southeastern Oregon. USDA For. Serv. Winward, A. H. Gen. Tech. Rep. PNW-80, 18 p. Pac. North- 1980. Taxonomy and ecology of sagebrush west For. and Range Exp. Stn., Portland, in Oregon. Agric. Exp. Stn. Stn. Bull. 642, Oreg. 15 p. Oreg. State Univ., Corvallis. Thomas, J. W., C. Maser, and J. E. Rodiek. Winward, A. H., andE. W. Tisdale. . 1979c. Edges. In Wildlife habitats in 1977. Taxonomy of the Artemisia managed rangelands: The Great Basin of tridentata complex in Idaho. For., Wildl., southeastern Oregon. USDA For. Serv. and Range Exp. Stn. Bull. 19, 15 p. Univ. Gen. Tech. Rep. PNW-85, 17 p. Pac. North- Idaho, Moscow. west For. and Range Exp. Stn., Portland, Oreg. Tisdale, E. W., M. Hironaka, and M. A. Fosberg. 1965. An area of pristine vegetation in Craters of the Moon National Monument, Idaho. Ecology 46(3):349-352. Tisdale, E. W., M. Hironaka, and M. A. Fosberg. 1969. The sagebrush region in Idaho, a problem in range resource management. Agric. Exp. Stn. Bull. 512, 15 p. Univ. Idaho, Moscow. Tueller, P. T. 1962. Plant succession on two Artemisia habitat types in southeastern Oregon. Ph. D. thesis. Oreg. State Univ., Corvallis. 249 p.

61 Appendix Common and Scientific Names of Animals

~ Common name Scientific name Symbol'

MAMMALS

Badger Taxidea taxus Beaver Castor canadensis Bighorn sheep Ovis canadensis Black-tailed jackrabbit Lepus californicus Bushy-tailed woodrat Neotoma cinerea Cattle Bos taurus Coyote Canis latrans Deer Odocoileus sp. Domestic sheep Ovis aries Domestic goat Capra sp. Gophers Thomomys sp. Ground squirrel Spermophilus sp. Hare Lepus sp. Kangaroo rat Dipodomys sp. Mule deer Odocoileus hemionus Pronghorn Antilocapra americana Rabbit Sylvilagus sp. Rocky Mountain elk Cervus elaphus Vole Lagurus sp.

BIRDS

Geese Branta sp. Horned lark Eremophila alpestris Lark sparrow Chondestes grammacus strigatus Sage grouse Centrocercus urophasianus Sage sparrow Amphispiza belli Swan Olor sp.

Common and Scientific Names and Symbols of Plants

TREES

Bit terc herr y Prunus emarginata PREM Chokecherry Prunus uirginiana PRVI Cottonwood Populus sp. POPUL Drummond willow Salix drummondiana SADR Fir Abies sp. ABIES Juniper Juniperus sp. JUNIP Ponderosa pine Pinus ponderosa PIP0 Quaking aspen Pop ul us tremu lo id es POTR Spruce Picea sp. PICEA Western juniper Juniperus occidentalis JUOC White fir Abies concolor ABCO

'After Garrison et al. (1976).

62 Appendix (continued)

SHRUBS

Antelope bitterbrush Purshia tridentata PUTR Basin big sagebrush Artemisia tridentata subsp. tridentata ARTRT Big sagebrush Artemisia tridentata ARTR Black greasewood Sarcobatus vermiculatus SAVE2 Black sagebrush Artemisia nova ARNO2 Bolander silver sagebrush Artemisia cana subsp. bolunderi ARCAB Broom snakeweed Gutierrezia samthrae GUSA Bud sagebrush Artemisia spinescens ARSP Bush cinquefoil Potentilla fruticosa POFR Carthamoid goldenweed Haplopappus carthamoides HACA Cleftleaf sagebrush Artemisia arbuscula subsp. thermopola ARART Coyote willow Salix exigua SAEX, Creeping Oregon grape Berberis repens BERE Curlleaf mountainmahogany Cercocarpus ledifolius ledifolius CELEL Desert rockspirea Holodiscus dumosus HODU Dwarf blueberry Vaccinium caespitosum VACA Dwarf threetip sagebrush Artemisia tripartita subsp. rupicola ARTRR Early low sagebrush Artemisia longiloba ARLO2 Goldenweed Haplopappus sp. HAPLO2 Granite gilia Leptodactylon pungens LEPU Gray horsebrush Tetradymia canescens TECA Gray rabbitbrush Chrysothamnus nauseosus CHNA Green rabbitbrush Chrysothamnus viscidiflorus CHVI Hop sage Grayia GRAY1 Horsebrush Tetradymia sp. TETRA Low sagebrush Artemisia arbuscula subsp. arbuscula ARARA Mountain big sagebrush Artemisia tridentata subsp. vaseyana ARTRV Mountain silver sagebrush Artemisia cana subsp. viscidula ARCAV Mountain snowberry Symphoriocarpus oreophilus SYOR Rabbitbrush Chrysothamnus sp. CHRYS3

Sagebrush ' Artemisia sp. ARTEM Saskatoon serviceberry Amelunchier alnifolia AMAL . Shadscale saltbush Atriplex confertifolia ATCO Silver sagebrush Artemisia cana ARCA Snow willow Salix nivalis SANI Spiny hopsage Grayia spinosa GRSP Squaw apple Pemph y 11 u m ra mos is s im u m PERA3 Stiff sagebrush Artemisia ngida ARRI Subalpine big sagebrush Artemisia tridentata form spiciformis ARTRS Threetip sagebrush Artemisia tripartita ARTR2 Wax currant Ribes cereum RICE Whiplash willow Salix caudata SACA Willow Salix sp. SALIX Wyoming big sagebrush Artemisia tridentata subsp. wyomingensis ARTRW

'After Garrison et al. (1976).

63 Appendix (continued)

Common name Scientific name Symbol'

GRASS AND GRASSLIKE PLANTS

Baltic rush Juncus balticus JUBA Bearded bluebunch wheatgrass Agropyron spicatum AGSP Big bluegrass Poa ampla POAM Blue wildrye Elymus glaucus ELGL Bottlebrush squirreltail Sitanion hys trix SIHY California brome Bromus carinatus BRCA Cheatgrass Brdmus tectorum BRTE Creeping bentgrass Agrostis palustris AGPA Crested wheatgrass Agropyron desertorum AGDE Cusick bluegrass Poa cusickii POCU Cutting wheatgrass Agropyron caninum AGCA Desert saltgrass Distichlis stricta DIST Douglas sedge Carex douglasii CAD0 Drummond rush Juncus drummondii JUDR Dunhead sedge Carex phaeocephala CAPH Dwarf squirreltail Sitanion hystrix var. hordeoides SIHYH Epapillous sedge Carex epap ill0 sa CAEP False-scirpus sedge Carex s c irp o id ea v a r . p s eu dos c irp o id ea CASCP Fescue Festuca sp. FESTU Giant wildrye Elymus cinereus ELCI Green fescue Festuca iiiridula FEVI H airgra s s Deschampsia sp. DESCH Idaho fescue Festuca idahoensis FEID Indian ricegrass Oryzopsis hymenoides ORHY Intermediate wheatgrass Agropyron intermed i u m AGIN2 Kentucky bluegrass Poa pratensis POPR Lemmon needlegrass Stipa lemmonii STLEZ Liddon sedge Carex petasata CAPE Longtongue mutton bluegrass Poa longiligula POLO Medusahead wildrye Elymus caput-medusae ELCAB Mountain hairgrass Desch a mps ia a trop u rp u rea DEAT Muhly Muhlenbergia richardsonis MURI Needleand thread Stipa comata STCO2 Needlegra s s Stipa sp. STIPA Nevada bluegrass Poa nevadensis PONE2 Northern meadow barley Hordeum b rac hy an therum HOBR One-spike oatgrass Dan thonia unispicata DAUN Oniongrass Melica sp. MELIC Ovalhead sedge Carex festivella CAFE Pacific fescue Festuca pacifica FEPA Pinegrass Calamagrostis rubescens CARU Prairie j unegrass Koeleria cristata KOCR Raynolds sedge Carex raynoldsii CARA Ross sedge Carex rossii CAR0 Rough fescue Festuca scabrella FESC Rush Juncus sp. JUNCU Sandberg bluegrass Poa sandbergii POSA3 Sedge Carex sp. CAREX Sheep fescue Festuca ovina FEOV Skyline bluegrass Poa epilis POEP Slenderbeak sedge Carex athrostachya CAAT Slender wheatgrass Agropyron trachycaulum AGTR

'After Garrison et al. (1976).

64 Appendix (continued)

Common name Scientific name Symbol'

Smallwing sedge Carex microptera CAM1 Smooth brome Bromus inennis BRIN Spike rush Eleocharis sp. ELEOC Spike trisetum Trisetum spicatum TRSP Spreading woodrush Luzula divaricata LUDI Subnigricans sedge Carex subnigncans CASU5 Swordleaf rush Juncus ensifolius JUEN Threadleaf sedge Carex filifolia CAFI Thurber needlegrass Stipa thurberiana STTH Tim0 thy Phleum pratense PHPR Western needlegrass Stipa occidentalis STOC Wheatgrass Agropyron sp. AGROP Wheeler bluegrass Poa nervosa PONE Winter bentgrass Agrostis scabra AGSC

FORBS

Alaska habenaria Habenaria unalascensis HAUN Alpine shootingstar Dodecatheon alpinum DOAL American bistort Polygonum bis tortoides POBI Annual agoseris Agoseris heterophylla AGHE Arrowleaf balsamroot Balsamorhiza sagittata BASA Ashy penstemon Penstemon humilis PEHU Aster Aster sp. ASTER Autum willowweed Epilo bium paniculatum EPPA Balsamroo t Balsamorhiza sp. BALSA Beckwith violet Viola beckwithii VIBE Bedstraw Galium sp. GALIU Beethistle Eryngium articulatum ERAR Bighead clover Trifolium macrocephalum TRMA Biscuitroot Lomatium sp. LOMAT Bloomer fleabane Erigeron bloomeri ERBL Bog-orc hid Habenaria sp. HABEN Buckwheat Eriogo n um caespi tos um ERCA Cinquefoil Potentilla sp. ' POTEN Clasping pepperweed Lepidium perfoliatum LEPE Columbia monkshood Aconitum columbianum ACCO Combleaf Polyctenium fremontii POFRB Common dandelion Taraxicum officianale TAOF Crag aster Aster scopulorum ASSC Crisped starwort Stellaria crispa STCR Daggerpod Phoenicaulis cheiranthoides PHCH Dandelion Taraxicum sp. TARAX Dwarf hesperochiron Hespemchimn pumilis HEPU Dwarf monkey flower Mimulus nanus MINA Elkslip marshmarigold Caltha leptosepala CALEZ False agoseris Microsens tmximoides M ITR False hellebore Veratrum sp. VERAT Fleabane Erigeron sp. ERIGE Gay penstemon Penstemon laetus PELA Gray hawksbeard Crepis intermedia CRIN Gray's biscuitroot Lomatium grayi LOGR Halogeton Halogeton glomemtus HAGL Hawksbeard Crepis sp. CREPI Heartleaf arnica Arnica cordifolia ARC0 'After Garrison et al. (1976).

65 Appendix (continued)

Common name Scientific name Symbol'

Holboell rockcress Ambis hoelboellii ARHO Hood's phlox Phlox hoodii PHHO Hooker balsamroot Balsamorhita hookeri BAHO Kellogg's knotweed Polygonum kelloggii POKE Lambstongue groundsel Senecio integerrimus SEIN Locoweed Astragalus sp. ASTRA Longleaf phlox Phlox longifolia PHLO Lousewort Pedicularis attollens PEAT Low pussytoes Antennaria dimorpha AND1 Lupine Lupinus sp. LUPIN Meadowrue Thalictrum sp. THALI Milkvetch Astragalus sp. ASTRA Modest buttercup Ranunculus verecundus RAVE Nevada biscuitroot Lomatium nevadenses LONE Newberry cinquefoil Potentilla newbenyi PONE3 Nineleaf biscuitroot Lomatium triternatum LOTR Nuttall violet Viola nuttallii VINU Oblongleaf bluebells Mertensia obligifolia MEOB Obscure milkvetch Astragalus obscurus ASOB Peavine Lathyrus sp. LATHY Phlox Phlox sp. PHLOX Pink microsteris Micros tens gracilis MIGR Plainleaf buttercup Ranunculus alismaefolius RAAL Prairiesmoke avens Geum trimrum GETR Primrose monkey flower Mimulus primuloides MIPR Pussy toes Antennaria sp. ANTEN Rockcress Arabis sp. ARAB1 Rocky Mountain parnassia Parnassia fimbriata PAFI Roseroot stonecrop Sedum roseum SERO Russian t his tle Salsola kali SAKA Scouler St. Johnswort Hypericum scouleri HYSC Short-leaved cinquefoil Potentilla brevifolia POBR Showy downingia Downingia elegans DOEL Sierra willowweed Epilobium brevistylum EPBR Skyrocket gilia Gilia aggregata GIAG Slenderbush eriogonum Eriogonum micro thecum ERMI Sorrel Rumex sp. RUMEX Spreading phlox Phlox diffusa PHDI Spring draba Draba vema DRVE2 Spur lupine Lupinus laxiflorus LULAB Starved milkvetch Astragalus miser ASMI Stenophyllus pussytoes Antennaria s tenophylla ANST Stinging nettle Urtica dioica URD1 Tapertip hawksbeard Crepis acuminata CRAC Thistle Cirsium sp. CIRSI Tiny mousetail Myosurus minimus MYMI2 Varileaf phacelia Phacelia he terophy lla PHHE Violet Viola sp. VIOLA Watson willowweed Epilobium watsonii EPWA Western hawkweed Hiemceum albertinum HIALB White bogorchid Habenaria dilatata HAD12 Wild buckwheat Eriogonum sp. ERIOG Woodlandstar Lithophragma bulbifera LIBU Woolly eriophyllum Eriophyllum lanatum ERLA Wyeth eriogonum Eriogonum hemcleoides ERHE Yarrow Achillea mille folium ACMI 'After Garrison et al. (1976).

66 WILDLIFE HABITATS IN MANAGED RANGELANDS - THE GREAT BASIN OF SOUTHEASTERN OREGON

Technical Editors

JACK WARD THOMAS, U.S. Department of Agriculture, Forest Service

CHRIS MASER, U.S. Department of the Interior, Bureau of Land Management

Title Now available Introduction

Plant Communities and Their ' Importance to Wildlife Gen. Tech. Rep. PNW-120 The Relationship of Terrestrial Vertebrates to the Plant Communities Native Trout Gen. Tech. Rep. PNW-84 Ferruginous Hawk Sage Grouse Pronghorn Mule Deer Bighorn Sheep Riparian Zones Gen. Tech. Rep. PNW-80 Edges Gen. Tech. Rep. PNW-85 Geomorphic and Edaphic Habitats Gen. Tech. Rep. PNW-99 Manmade Habitats Gen. Tech. Rep. PNW-86 Management Practices and Options The Forest Service of the U.S. Department of Agriculture is dedicated to the principle of multiple use management of the Nation’s forest resources for sustained yields of wood, water, forage, wildlife, and recreation. Through forestry research, cooperation with the States and private forest owners, and management of the National Forests and National Grasslands, it strives - as directed by Congress - to provide increasingly greater service to a growing Nation. The U.S. Department of Agriculture is an Equal Opportunity Employer. Applicants for all Department programs will be given equal consideration without regard to age, race, color, sex, religion, or national origin.

GPO 982-914